ub 


A   REVISION    OF    THE    ADULT    TAPEWORMS    OF   HARES 

AND  RABBITS.1 


By  OH.  WARDELL/STILES,  Ph.  D., 

Honorary  Custodian  of  the  Helminthologieal  Collection,  U.  S.  National  Museum. 


A  DISCOVERY  by  Cooper  Curtice  in  1887  regarding  certain  young 
stages  of  rabbit  tapeworms,  observations  which  I  was  able  to  confirm 
and  extend  in  1894.  immediately  brought  the  leporine  cestodes  into 
a  very  important  position ,  viewed  from  an  economic  as  well  as  from  a 
scientific  standpoint,  for  Curtice's  observation  at  first  sight  seemed  to 
offer  a  guide  to  solving  the  puzzle  as  to  the  larval  stage  of  the  tape- 
worms of  cattle,  sheep,  horses,  and  certain  other  animals. 

Curtice  observed  in  the  intestine  of  rabbits  certain  young  cestodes 
which  bore  minute  hooks  upon  the  rostellum,  and  older  stages  were 
foTTiid  which  had  lost  these  hooks.  He  determined  the  adult  cestodes 
found  in  these  rabbits  as  Tcenia  pectinata  andflooked  upon  the  armed 
forms  as  the  young  of  this  species.  .i 

With  these  observations  before  us,  it  lookedjas  if  some  very  radical 
changes  would  be  made  in  the  near  future  in  Jhe  classification  of  the 
tapeworms  of  the  larger  domesticated  animals;  it  further  seemed  prob- 
able that  the  young  stage  of  the  tapeworms  of  cattle,  sheep,  and  horses, 
would  eventually  be  found  to  be  an  armed  cysticercoid. 

In  a  paper2  in  1894  I  called  attention  to  this  bearing  of  the  question 
and  extended  Curtice's  observations.  I  found  that  the  suckers3  as 
well  as  the  rostellum  were  armed,  and  that  the  head  bore  a  remarkable 
resemblance  to  the  head  of  Davainea.  This  complicated  the  question 
in  so  far  as  to  render  necessary  the  consideration  of  two  further  pos- 
sibilities— i.  e.,  were  these  young  cestodes  the  early  stages  of  avian 


1  At  the  request  of  the  author,  it  is  here  stated  that  he  is  not  responsible  for  the 
insertion  of  commas  between  the  names  and  the  authorities,  and  for  the  absence  of 
capitals  at  the  beginning  of  specific  names  derived  from  personal  names.  These 
changes  have  been  made  in  order  to  bring  his  paper  into  conformity  with  the  usages 
at  present  followed  in  the  U.  S.  National  Museum. — EDITOR. 

-Notes  sur  les  Parasites — 31 :  line  phase  precoce  des  Te"nias  du  Lapin  (Notice  pre"- 
liminaire),  Bull.  Soc.  zool.  France,  1894,  XIX,  pp.  163-165. 

3Some  old  sketches  of  Curtice's  show  that  ho  also  observed  the  hooks  upon  the 
suckers. 


PROCEEDINGS  U.  S.  NATIONAL  MUSEUM,  VOL.  XIX—  No.  1105. 

*  145 

Proc.  N.  M.  vol.  xix  -  10  ^*X2<£Z^ 


M363S29 


146  TAPEWORMS  OF  HAUES  AND  RABBITS— STILES.         VOL.XIX. 

cestodes  Avhich  had  accidentally  gained  access  to  rabbits,  or  did  there 
exist  in  American  rabbits  an  adult  cestode  belonging  to  the  genus 
Davaineaf 

The  relation  of  the  subject  to  the  tapeworms  of  cattle,  sheep,  and 
horses  made  it  absolutely  necessary  from  an  economic  standpoint,  and 
the  relation  of  these  forms  to  the  adults  in  rabbits  made  it  desirable 
from  a  scientific  standpoint,  to  immediately  revise  the  adult  cestodes 
found  in  rabbits.  This  work  was  accordingly  begun,  and  in  1895  my 
paper1  was  published  announcing  the  finding  of  a  double-pored  cestode 
with  occasional  single  pores,  and  the  occurrence  in  American  rabbits 
of  a  single-pored  cestode  with  a  Davainea-lik&  uterus. 

This  paper  practically  disposed  of  the  young  armed  cestode  of  rabbits, 
for  after  reading  it  the  natural  conclusion  of  every  helmiuthologist 
would  be  that  this  parasite  was  probably  a  Davainea  and  that  we  were 
no  nearer  the  solution  of  the  question  as  to  the  young  form  of  Moniezia 
than  we  were  before. 

The  present  paper  is  the  result  of  the  revisional  study  mentioned 
above.  In  it  I  finally  dispose  of  the  armed  form  mentioned  in  1894, 
and  at  the  same  time  I  am  obliged  to  complicate  the  question  as  to  the 
larval  stage  of  Moniezia  still  further  by  presenting  an  unarmed  young 
cestode  in  rabbits.  I  further  give  an  anatomical  systematic  revision  of 
the  adult  leporine  tapeworms,  together  with  their  generic  relationship 
to  the  cestodes  of  cattle,  sheep,  and  horses. 

I  desire  to  call  particular  attention  to  the  young  unarmed  cestode  de- 
scribed on  p.  201.  Had  any  worker  found  this  parasite  in  an  insect, 
worm,  or  snail  in  a  locality  where  cattle  and  sheep  are  infested  with 
Moniezia  expansa,  he  would  hardly  have  hesitated  to  announce  the  dis- 
covery of  the  intermediate  host  of  this  important  parasite.  The  find- 
ing in  one  host  of  a  larval  form  whose  head  resembles  a  certain  adult 
in  another  host  does  not,  however,  scientifically  establish  the  life  history 
of  that  particular  parasite, 

I  can  now  prophesy  confidently  that  it  will  be  almost,  if  not  entirely, 
impossible  to  distinguish  the  larval  stage  of  M.  expansa  from  that  of  a 
dozen  or  so  of  other  tapeworms,  and  on  account  of  the  great  economic 
importance  of  this  question  I  caution  against  any  too  early  and  too 
ungrounded  announcement  of  the  source  of  infection  of  cattle  and  sheep 
by  this  species.  The  only  work  upon  this  subject  which  will  be  worthy 
of  full  credence  is  experimental  feeding. 

Within  recent  years  it  has  been  customary  to  associate  the  larvae 
found  parasitic  in  some  animals  with  the  adults  parasitic  in  other  hosts 
simply  because  of  a  similarity  of  the  heads  and  hooks.  This  is  par- 
ticularly the  case  with  the  avian  tapeworms.  I  feel  it  necessary  to  enter 
a  protest  against  carrying  these  generalizations  too  far,  for  at  present, 
when  so  many  of  the  adult  avian  parasites  are  so  incompletely  described 

1  Notes  on  Parasites— 36:  A  double-pored  cestode,  with  occasional  single  pores, 
Centralbl.  f.  Bakter.  u.  Parasitenk.  .  1  Abt..  XVII,  13-14,  pp.  457-459. 


PROCEEDINGS  OF  TJfj:  NATIOXAL  MUSEUM.  147 


that  they  can  scarcely  be  recognized,  it  can  hardly  be  expected  that  the 
larvie  can  be  determined  with  certainty.  The  work  by  Mrazek,  Moniez, 
Ilamann,  von  Linstow,  and  others  in  describing  these  larval  forms  has 
been  most  valuable ;  but  authors  have,  I  believe,  generalized  too  much 
upon  these  observations.1 

Acknowledgments. — I  am  indebted  to  the  following  gentlemen  for  fur- 
nishing me  with  specimens  for  study  in  preparing  this  revision : 

Prof.  Eaphael  Blanchard,  of  Paris:  A  portion  of  Baird's  type  of 
Tcenia  goezei;  specimens  of  Anoplocephala  wimerosa  and  Cittotcenia 
leuckarti.  Dr.  Gustav  Brandes,  of  Halle:  Original  cotypes  of  Eiehm's 
Dipylidium  pectinatum,  D.  leuckarti,  and  D.  latissimum.  Geheimrath 
Karl  Mobius  and  Dr.  A.  Collin,  of  Berlin:  Fragment  of  Kudolphi's 
Tcenia pectinata.  Prof.  Herbert  Osborn,  of  Ames,  Iowa:  Specimens  of 
Cittotamia  prcecoquis.  Prof.  M.  J.  Elrod,  of  Bloomington,  Illinois :  Speci- 
mens of  Bertia  americana.  Mr.  J.  H.  Tallichet,  of  Austin,  Texas :  Speci- 
mens of  Davainea  salmoni  and  Cittotcenia  variabilis.  Dr.  A.  K.  Fisher, 
Division  of  Mammalogy  and  Ornithology,  United  States  Department 
of  Agriculture:  Specimens  of  Davainea  rctractilis.  Mr.  Eobert  A. 
Mills,  of  Chuluota,  Florida:  Specimens  of  Cittotcenia  variabilis  imbri~ 
cata.  Dr.  Austin  Peters,  of  Boston,  Massachusetts:  Specimens  of  C. 
variabilis, 

The  other  material2  used  belonged  to  collection  of  Bureau  of  Animal 
Industry  (collected  by  Curtice,  Hassall,  Stiles)  5  collection  of  the  United 
States  National  Museum;  collection  of  Leidy  (University  of  Pennsyl- 
vania), and  collection  of  Stiles  (United  States  National  Museum). 

Although  this  article  treats  primarily  of  leporine  cestodes,  it  has 
been  found  necessary  to  include  several  forms  from  other  hosts  for 
comparison. 

To  my  assistant,  Albert  Hassall,  I  am  indebted  for  the  entire  technique 
connected  with  the  specimens  used,  and  also  for  the  preparation  of  the 
bibliography,  and  to  Mr.  W.  S.  D.  Haines,  artist  of  the  Bureau  of  Ani- 
mal Industry,  Department  of  Agriculture,  for  preparing  the  plates. 

Unfortunately  much  of  the  material   at  my  disposal  was  poorly 

1  For  a  detailed  discussion  of  this  subject  cf.  Stiles :  Keport  upon  the  Present  Knowl- 
edge of  the  Tapeworms  of  Poultry,  Bull.  12,  Bureau  of  Animal  Industry,  U.  S.  Depart- 
ment of  Agriculture,  1896,  pp.  7-73,  pis.  i-xxi. 

2  Since  issuing  my  preliminary  note  on  this  group,  I  have  received  the  following 
specimens  from  the  gentlemen  mentioned : 

Prof.  A.  Moniez,  of  Lille:  His  entire  collection  of  auoplocephaline  cestodes, 
including  the  type  specimen  of  Tcenia  wimerosa  and  undetermined  specimens  of 
Cittoicenia  denticulata  and  C.  pectinata.  Geheimrath  Rudolf  Leuckart,  of  Leipzig: 
cotypes  of  Riehm's  Tcenia  rhopalocepliala  and  T.  rhopaliocephala  and  one  of  Meyner's 
originals  of  Tcenia  (Bertia)  mucronata.  Dr.  Victor  A.  Norgaard,  of  Alice,  Texas: 
Specimens  of  Davainea  salmoni.  Dr.  O.  Aron  Linstow,  of  Gottingen :  Specimen  of 
Cittotcenia  pectinata.  Dr.  von  Marenzeller,  of  Vienna :  Several  of  Riehm's  cotypes. 
Dr.  Collin,  of  Berlin  :  Rudolphi's  cotypes  of  Tcenia  denticulata. 

The  manuscript  of  this  article  was  delayed  in  order  that  these  specimens  could 
be  studied  and  the  results  incorporated  in  the  revision. 


148  TAPEWORMS  OF  HARES  AXD  RABBITS— STILES.         VOL.XIX. 

preserved,  so  tliat  a  number  of  points  I  have  been  obliged  to  leave  for 
further  study.  The  present  study,  however,  reduces  the  known  leporine 
forms  (Andrya  and  Bertla  excepted)  to  a  comparatively  satisfactory 
system. 

It  must  be  held  in  mind  that  every  classification  proposed  for  the 
Cestoda  for  some  years  to  come  is  experimental  and  provisional,  for 
helminthology  is  not  yet  so  far  advanced  that  we  can  tell  with  any 
degree  of  certainty  what  characters  or  combinations  of  characters 
should  be  looked  upon  as  of  family,  subfamily,  generic,  subgeneric, 
specific,  and  varietal  value,  and  what  characters  should  be  attributed 
to  host-influence. 

Date  of  American  species. — "  Notes  on  Parasites — 38 :  Preliminary  note 
to  <A  Revision  of  the  Adult  Leporine  Oestodes,' ni  established  August 
28,  1895,  as  the  date  of  the  new  specific  names  used  in  this  paper. 

Important  notice  to  helminthologists. — Owing  to  the  fact  that  Hassall 
and  I  have  been  using  material  from  different  helminth ological  collec- 
tions, more  particularly  from  the  collection  of  the  Bureau  of  Animal 
Industry,  collection  of  the  United  States  National  Museum,  collection 
of  Hassall,  collection  of  Stiles,  and  collection  of  Leidy  ( University  of 
Pennsylvania)  bearing  parallel  current  numbers,  some  confusion  has 
arisen  because  the  same  number  sometimes  appears  in  all  five  collec- 
tions. These  five  collections  were  commenced  independently,  and  as 
their  union  (temporary  or  permanent)  was  not  foreseen,  this  duplication 
of  numbers  could  not  be  avoided. 

To  avoid  such  confusion  in  the  future  as  far  as  possible,  the  collection 
of  the  Bureau  of  Animal  Industry  and  collection  of  Stiles  will  be  cov- 
ered into  the  United  States  National  Museum,  and  receive  the  current 
numbers  of  the  helmiuthological  collection  of  the  Museum. 

The  numbers  which  have  been  published  will  not  be  affected  by  this 
change,  so  far  as  can  now  be  foreseen.  The  numbers  of  a  few  of  the 
specimens  Avhich  have  been  distributed,  however,  will  in  all  probability 
be  changed.  Records  of  these  specimens  have  been  kept  in  the  Zoolog- 
ical Laboratory,  Bureau  of  Animal  Industry,  and  new  labels  bearing 
the  new  numbers  will  be  written  and  mailed  to  specialists  and  museums 
possessing  such  specimens  as  are  affected  by  the  renumbering  of  the 
collections. 

The  Leidy  collection,  as  I  have  stated  in  another  publication,  is  depos- 
ited with  me  only  temporarily  and  will  be  eventually  returned  to  the 
University  of  Pennsylvania.  Duplicates,  however,  will  be  kept  with 
the  United  States  National  Museum. 


Family 

Diagnosis. — Cestoda  with  distinctly  segmented  strobila;   head  pro- 
vided with  four  cup  shaped   suckers;   rostellum   well  developed,  or 

1  Vet.  Mag.,  II,  June,  1895,  pp.  341-346,  issued  Aug.  28, 1895. 


NO.  1 105.  PR  0  CEEDINGS  OF  THE  NA  TTO  KA  L  M  VSE  UM.  149 


rudimentary  and  not  always  evident  (?  or  entirely  absent)  5  genital 
pores  generally  lateral ;  uterus  without  special  pore. 

Type  genus. — Tcenia,  Linnaeus,  1758. 

All  of  the  leporine  cestodes  known  at  present  belong  to  the  family 
Tfieniidae,  in  which  most  authors  place  indiscriminately  all  tapeworms 
provided  with  four  cup-like  suckers.  Were  this  family  subjected  to 
revision  by  any  set  of  zoologists  except  helminthologists,  it  would 
undoubtedly  soon  be  raised  to  superfamily  rank  at  least,  and  divided 
into  several  families,  subfamilies,  and  numerous  genera.  Most  hel- 
minthologists, however,  are  very  conservative  in  proposing  genera, 
even  when  very  prolific  in  establishing  species!  A  superfamily  Treni- 
oidea3  will,  I  believe,  soon  be  established,  but  this  step  it  will  be  well 
to  postpone  until  the  cestodes  of  birds  are  brought  into  better  order. 


ANALYTICAL   KEY   TO    THE    GENERA   OF   ADULT   TAPEWORMS   FOUND   IN   LEPUS.* 


1.  Head  unarmed;  genital  pores  single  or  double;  ova  with  pyriform  body. 

Anoploceplialince  2 

Head  arrned  with,  numerous  hooks  on  the  rostelluin  and  suckers;  genital  pores 
single;  ova  without  pyriform  body Davainea  (p.  194.) 

2.  Genital  pores  double;  single  pores  a  rare  exception  and  then  in  strobilse  contain- 

ing double  pores Cittotcenia  (p.  170.) 

Genital  pores  single 3 

3.  Genital  pores  unilateral;  testicles  unilateral  or  nearly  so,  in  portion  of  median 

field  opposite  pores ;  uterus  transverse  with  proximal  and  distal  egg  pouches; 

no  separate  and  distinct  prosfcatic  gland  present Anoplocepliala  (p.  150.) 

Genital  pores  irregularly  alternate;  testicles  extend  across  the  median  field  to  or 
beyond  the  ovary ;  uterus  appears  as  a  network  or  as  a  transverse  tube  with 
egg  pouches 4 

4.  Separate  and  distinct  prostata  wanting ;  uterus  a  transverse  tube  with  proximal 

and  distal  egg  pouches Bertia  (p.  160.) 

Separate  and  distinct  prostata  present ;  uterus  reticulate ;  genital  pores  show  a 
marked  tendency  to  uiiilaterality Andrya  (p.  154.) 

Subfamily  ^IVOIPLOCEFHAJLIIS^E;,  R.  BlaneharcU  1891. 

Diagnosis. — Tseniidre  with  unarmed  head;  genital  pores  lateral, 
single  or  double;  segments  nearly  always  broader  than  long;  uterus 
transverse  and  tubular,  or  reticulate;  ventral  canals  always  well  devel- 
oped; dorsal  canals  generally  less  developed;  ova  generally  with  pyri- 
form body;  calcareous  bodies  present  or  absent. 

Type  genus. — Anoplocepliala^  E.  Blanchard,  1848. 

Kiehm2  in  1881  placed  the  three  double -pored  rabbit  cestodes  known 
to  him  in  the  genera  DipyUdium,  R.  Leuckart,  and  Cittotcenia^  Riehm, 
and  the  single-pored  forms  in  Tcenia;  in  his  second3  paper  he  placed 


!See  also  p.  213. 


1  See  also  p.  213. 

2Zeitschr.  ges.  Naturw.,  3  ser.,  VI,  p.  200. 

3Studien  an  Cestoden,  Zeit.  ges.  Naturw.,  3  ser.,  VI,  pp.  545-610. 


l,r)0  TAPEWORMS  OF  HARES  AXD  RABBITS— STILES.         voi.xix. 

all  the  double-pored  forms  in  Dipylidium.  R.  Blanchard1  in  1891 
placed  the  double-pored  forms  in  Moniezia,  the  single -pored  forms  in 
Anoplocepliala;  Stiles 2  in  1893  separated  the  double-pored  forms  from 
Moniezia,  but  did  not  establish  any  genus  for  them;  Railliet3  in  1893 
created  the  genus  Ctenotcenia  for  the  double-pored  forms  and  An  dry  a 
for  the  single-pored  forms. 

My  studies  now  lead  me  to  adopt  Cittotcenia  for  the  double-pored 
leporine  cestodes,  and  to  divide  the  unarmed  single-pored  forms 
between  the  genera  Anoplocepliala^  Andrya,  and  Bertia. 

Genus    ANOPLOCEPHALA,     E.      Blanehard,    1848    (nee 

Stal  187O). 

1848,  Anoplocephala,  E.  BLANCHARD,  Ann.  des  Sci.  nat.,  3  ser.,  Zool.,  X,  p.  344.     Type 

by  elimination,  Tcenia  equina  perfoliata,  Goeze,  1782. 
1871,  Plagiotccnia,*  PETERS,  Proc.  Zool.  Soc.  Lond  ,  p.  146.      Type,  Tcenia  gigantea, 

Peters,  1856. 
1891,  " Plagotcenia,  PETERS,  1871  "(as  synonym)  in  R.  BLANCHARD,  Me~m.  Soc.  zool. 

France,  IV,  p.  446. 

Diagnosis. — Anoplocephalinae  with  segments  broader  than  long. 
One  set  each  of  male  and  female  organs  in  each  segment;  genital  pores 
unilateral  (dextral5) ;  testicles  in  the  aporose,  ovary  in  the  pore  side  of 
the  median  field.  Uterus  a  transverse  tube  with  proximal  and  distal 
pouches.  Dorsal  canals  lie  dorsal  or  lateral  of  ventral  canal.  Genital 
canals  cross  the  longitudinal  canals  and  nerves  dorsally.  Calcareous 
bodies  (always?)  absent  from  parenchyma.  Eggs  with  well-developed 
pyriform  body.  Hosts:  Perissodactyla,  Kodentia. 

Type.— A.  perfoliata  (Goeze,  1872),  E.  Blanehard,  1848. 

1  Sur  les  Helminthes  des  Primates  Anthropoides,  Me"m.  Soc.  zool.  France,  IV,  pp. 
186-196.     Notices  helminthologiques :  7. — Cestodes  du  groupe  des  Anoplocephalime 
R.  Blanchard,  1891,  Me"m.  Soc.  zool.  France,  IV,  pp.  443-450. 

2  A  Revision  of  the  Adult  Cestodes  of  Cattle,  Sheep,  and  Allied  Animals,  Bull.  IV, 
Bur.  An.  Ind.,  U.  S.  Dept.  Agric. 

3Trait<5  Zool.  me'd.  et  agric.,  I. 

4  Max  Braiin  (1894,  Verrnes,  36-37  Lieferung,  p.  1138)  erroneously  gives  the  date  of 
this  genus  as  1870;  R.  Blanehard  spells  the  generic  name  Plagotcenia.  A  typograph- 
ical error  in  R.  Blanehard,  1891,  p.  446,  gives  Anoplocepliala  the  date  of  1868,  while 
Brauu  (loc.  cit.,  p.  1133)  dates  the  genus  1847.  Errors  of  this  kind  naturally  creep 
into  every  author's  writing.  In  this  connection  I  would  call  attention  to  the  list  of 
Cestoda  given  hy  Braun  (loc.  cit.,  pp.  1133-1145).  While  this  list  is  extremely  valua- 
ble in  tracing  the  different  species,  it  must  not  be  looked  upon  as  complete;  further- 
more, care  must  be  exercised  in  using  the  dates  of  species  given  by  Brauu.  Rudolphi's 
species  of  1810  are  erroneously  given  as  1808 ;  Krabbe's  species  of  1869  are  erroneously 
given  as  1870.  A  number  of  other  species  are  also  incorrectly  dated. 

5 In  A.wimerosa  the  pores  are  evidently  dextral,  but  in  the  other  species  I  am 
unable  to  determine  whether  they  are  dextral  or  sinistral,  either  from  the  figures  or 
descriptions  given  by  various  workers.  In  one  of  my  preparations  of  A.  mamiUana 
they  are  certainly  sinistral;  in  another  preparation  they  are  apparently  dextral. 
Professor  Zschokke  has  reexamined  his  preparations  of  A.  mamiUana  at  my  request 
and  writes  me  that  he  finds  the  pores  dextral,  which  agrees  with  the  topography 
found  upon  a  preparation  by  Railliet  very  recently  sent  to  me. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  151 


E.  Blanchard  in  1891  admitted  the  following  species  to  tins  genus: 


A.perfollata1  (Goeze,  1782). 
A.  mamillana  (Mehlis,  1831). 
A.  plicata  (Zeder,  1800). 
A.  transversaria  (Krabbe,  1879). 
A.  ivimerosa  (Moniez,  1880). 
A.  globiceps2  (Diesing,  1856). 


A.  blanchardi,  Moniez,  1891. 

.1.  zebrce  (Rudolphi,  1810). 

A.  hijracis  (Rudolphi,  1810). 

A.  gifjantea  (Peters,  1856). 

Tcenia  rhopaloeephala,  Riehui,  1881. 

T.  rhopaliocephala,  Riehm,  1881. 


Two  of  these  species,  Tcenia  rhopaloeephala  and  T.  rhopaliocephala, 
I  refer  with  Railliet  to  the  genus  Andrya  (vide,  p.  154).  Of  the 
other  species,  A.  wimerosa  is  the  only  one  which  occurs  in  rabbits. 
This  form  is  very  closely  allied  to  A.  mamillana  of  the  horse  and 
A.  transversaria  of  the  marmot,  which  are  here  introduced  for  com- 
parison. Several  of  the  remaining  forms  require  further  study  before 
their  generic  position  can  be  looked  upon  as  fully  established.  Setti 
(1893)  refers  Arhynchotwnia  critica  Pagenstecher,  1877,  and  Tcenia 
ragazzii  Setti,  1891,  from  Hyrax  to  this  genus. 

ANOPLOCEPHALA  WIMEROSA  (Moniez,  1880),  R.  Blanchard,  1891. 

(Plato  V,  figs.  1-7.) 

1880,3  Tcenia  wimerosa,  MONIEZ,  Bull,  scientif.  du  Depart,  dn  Nord,  2  ser.,  3  ann.,  no.  6, 

Juin,  pp.  240-242. 
1891,  Anoplocephala  ivimerosa  (MONIEZ,  1880),  R.  BLANCHARD,  Mem.  Soc.  zool.  France, 

IV,  p.  187,  p.  449. 
1893,  Andrya  wimerosa  (MoNiE/,  1880),  RAILLIET,  Traite"  de  Zool.  med.  et  agric.,  I,  p. 

283. 

Moniez,  in  1880,  described  as  T.  wimerosa  a  cestode  which  he  found 
in  Lepus  cunlculus  at  Wimereux.  His  description  reads  as  follows : 

[p.  241.]  Le  Tcenia  Wimerosa  appartient  au  type  du  Tccnia  expansa.  Observe"  a,  Trail 
mi  ou  sous  de  faibles  grossissements,  cette  espece  qui  atteint  a  peine  un  centimetre  de 
long  sur  une  largeur  de  un  millimetre  et  demi,  se  pre"sente  avec  un  corps  e"pais,  forme" 
d'une  dizaine  d'anneaux  seulemeut.  La  tete  est  grosse,  les  ventouses  6cartees,  il 
n'y  a  ni  bulbe  ni  crochets,  le  con  est  nul.  Les  anneaux  s'accusent  d'abord  par  des 
plis  accentues;  leur  rebord  inf<5rieur  tres  saillant  est  arrondi  et  orn6  d'une  serie  de 
cils  616gamment  disposes;  leur  aspect  rappelle  celui  des  cils  des  ventouses  dont  j'ai 
parle"  ailleurs  a  propos  de  la  Ligule. 

L'appareil  genital  n'est  pas  double  dans  cette  espece  comme  cLez  beaucoup 
d'Inermes  et,  par  une  autre  particularity  tous  les  anneaux  le  portent  du  ineme  cote. 
En  meme  temps,  1'ouverture  gcnitale  d^bouche  au  rebord  infdrieur  de  Tanneau,  bien 

1  The  tapeworms  of  the  horse  should  be  subjected  to  an  anatomical  revision,  and  at 
the  same  time  their  specific  names  should  be  correctly  established.  I  refrain  from 
attempting  to  straighten  out  these  names  at  present,  as  this  can  be  done  satis- 
factorily only  when  one  has  a  good  line  of  specimens  before  him.  The  tapeworms 
of  horses  are  unfortunately  very  poorly  represented  in  the  Bureau  of  Animal  Indus- 
try collection. 

2Llihe,  1895  B,  pp.  202-205,  has  recently  reexamined  Diesing's  type  material  of  this 
species,  and  has  shown  it  to  be  an  unquestionable  Anoplocephala. 

3 The  citations  immediately  following  the  specific  names  include  only  those  articles 
which  have  a  direct  bearing  on  the  synonymy  s.  st. ;  other  references  are  cited  in  the 
text  by  date  of  publication.  (See  Bibliography,  p.  222.) 


152  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 

que  la  pocbe  pe~niale  soit  situee  exactemeiit  en  son  milieu ;  le  pe"nis,  qui  est  trrs  long, 
decrit  done  pour  sortir  une  courbe  accentuee.  La  saillie  que  ce  dernier  organ e  peut 
faire  en  dehors  est  considerable. 

Le  de"veloppement  de  la  poche  peniale  iniprime  des  modifications  particulieres  a  la 
forme  de  cet  animal.  La  poche  pe~niale  occupe  d'abord  la  plus  grande  partie  de 
1'anneau;  par  suite  de  la  rapidite"  de  son  developpement,  qui  marche  beaucoup  plus 
vite  que  celui  du  reste  de  1'anneau,  elle  forme,  sur  le  bord,  une  saillie  tres  forte  qui 
e'efface  a  mesure  que  1'anneau  grandit.  La  syme'trie  se  trouve  ainsi  rctablie,  mais 
le  developpement  des  ceufs  fait  bientot  disparaltre  completement  1'dnorme  poclie 
peniale. 

Les  forts  "grossissements  permettent  de  voir  les  oeufs ;  dont  les  caracteres  sont 
ceux  des  oeufs  des  Ttenias  inermes  [p.  242]  vrais;  leur  appareil  pyriforme  est  tres 
developpe".  Les  muscles  longitudinaux  sont  gros  et  forment  une  zone  continue  et 
peu  e"paisse.  La  z6ne  de  proliferation  est  tres  dtendue  et  sous-jacente  aux  ventouses, 

Blancliard  in  1891  found  the  same  form  in  Lepus  variabilis  and 
determined  it  as  an  Anoplocephala. 

Eailliet  in  1893  transferred  it  to  his  genus  Andrya. 

Through  the  kindness  of  Blanchard,  I  obtained  a  number  of  his  speci- 
mens upon  which  the  following  details  are  based : 

The  worms  attain  10  mm.  in  length  by  2  mm.  in  breadth.  The  head 
is  unarmed,  nearly  square  when  viewed  en  face,  measuring  about  0.78 
to  0.88  mm.  broad ;  it  sits  like  a  knob  on  the  end  of  the  strobila,  from 
which  it  is  sharply  defined.  No  rostellum  was  visible.  The  four  suck- 
ers are  powerful,  having  a  diameter  of  nearly  0.4  mm.  Neck  absent 
segmentation  beginning  immediately  back  of  the  head;  as  many  as  28 
segments  are  present  in  some  individuals.  The  anterior  segments  are 
much  broader  than  long  and  somewhat  asymmetrical;  the  posterior 
segments  may  measure  2.24  mm.  broad  by  0.8  mm.  long;  in  some  cases 
they  become  nearly  as  long  as  broad.  The  genital  pores  are  unilateral 
and  dextral. 

The  reproductive  glands  are  confined  to  the  anterior  third  of  the 
worm,  while  the  posterior  two-thirds  are  occupied  by  the  uterus.  The 
genital  pore  becomes  almost  or  quite  obliterated  upon  the  atrophy 
of  the  glands.  The  testicles  are  confined  entirely  to  the  aporose  side 
of  the  segment,  as  is  the  case  with  Anoplocephala  mamillana  and 
A.  transversaria. 

This  parasite  is  very  closely  allied  to  A.  mamillana  of  the  horse. 
From  the  descriptions  of  former  authors  and  from  my  own  studies,  I 
propose  the  following  as  a  revised  specific  diagnosis: 

Diagnosis. — Anoplocephala  wimerosa  (Moniez,  1880),  E.  Blancliard, 
1891:  Strobila  attains  10  mm.  in  length  by  1.5  to  2.25  mm.  in  breadth, 
and  contains  from  10  to  28  segments  which  are  always  broader  than 
long;  proximal  segments  often  asymmetrical  in  outline.  Distal  seg- 
ments attain  about  2.25  mm.  broad  by  0.8  mm.  long,  rarely  becoming 
nearly  as  long  as  broad.  Head  unarmed,  nearly  cuboid,  measuring  0.7 
to  0.88  mm.,  and  sits  like  a  knob  on  the  end  of  the  strobila;  rostellum 
not  observed;  suckers  0.4  mm.  in  diameter,  prominent,  rounded,  open- 
ing diagonally  forward ;  posterior  lobes  absent.  Keck  absent.  Gem 
tal  pores  unilateral,  dextral.  The  sexual  glands  are  confined  to  the 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  153 

proximal  third  of  the  worm,  the  uterus  occupying  the  distal  two-thirds. 
Male  organs:  Testicles  about  15  to  30  in  a  segment,  left  of  the  median 
line 5  they  appear  in  the  first  or  second  segment,  and  atrophy  by  the 
fifteenth;  cirrus-pouch  dorsal  of  vagina,  attains  0.48  mm.  in  length, 
crossing  the  longitudinal  canals  dorsally.  Female  organs:  Ovary 
appears  in  earliest  segments  in  about  the  median  line,  and  atrophies 
earlier  than  the  testicles;  receptaculum  seminis  elongate;  uterus  ven- 
tral, transverse  with  distal  and  proximal  pouches.  Ova  52  JJL  iu  diame- 
ter, pyriform  body  12  //,  horns  crossed.  Dorsal  canal  dorsal  to  lateral 
of  ventral  canal.  Genital  canals  pass  dorsally  of  longitudinal  canals 
and  nerves. 

Jlosts. — European  Eabbit  (Lepus  cunlculus]  by  Moniez;  Mountain 
Hare  (L.  variabilis)  by  E.  Blanchard. 

Type. — No.  1452,  U.S.N.M.,  belongs  to  collection  of  Moniez.  Typi- 
cal specimens  in  collection  of  E.  Blanchard  (Paris);1  collection  of 
Bureau  of  Animal  Industry;1  collection  of  Stiles  (U.S.N.M.);1  Nos. 
112,  1353,  1358,  1359,  1360,  1361,  U.S.N.M.;1  collection  of  Hassall;1 
collection  of  Leidy  (University  of  Pennsylvania) ; T  collection  of  Har- 
vard University;1  collection  of  II.  B.  Ward. 

Geographical  distribution. — France:  Wimereux  (by  Moniez);  Brian- 
§on,  by  E.  Blanchard. 

ANOPLOCEPHALA   MAMILLANA  (Mehlis,  1831),  R.  Blanchard,  1891. 
(Plates  V,  fig.  8;  VI,  figs.  1-3.) 

1831,  Tcenia  mamillana,  MEHLIS,  Gurlt's  Lehrbuch  d.  path.  Anat.  d.  Haussaugetliiere, 

I,  p.  380,  pi.  ix,  figs.  7, 11. 
1891,  Anoplocepliala  mamillana  (MEHLIS,  1831),  R.  BLANCHARD,  Me'm.  Soc.  zool.  France, 

IV,  p.  187. 

Diagnosis. — Anoplocephala  mamillana  (Mehlis,  1831),  E.  Blanchard, 
1891:  Strobila  attains  6  to  30  mm.  in  length  by  4  to  6  mm.  in  breadth. 
Head  unarmed,  0.7  to  0.8  mm.  broad  by  0.5  mm.  long;  suckers  very 
muscular,  elliptical  with  elongate  openings;  posterior  lobes  absent. 
Neck  absent.  Thirty-five  to  fifty-two  segments  present,  the  distal  4  to 
8  completely  filled  with  ova;  sexually  active  segments  3  to  5  mm. 
broad  by  0.3  to  0.6  mm.  long;  posterior  segments  may  attain  2  mm.  in 
length.  Genital  pores  unilateral  in  posterior  half  of  lateral  margin. 
Male  organs:  Testicles  confined  to  aporose  side  of  segment, appearing 
in  the  first  segments,  and  atrophying  by  the  seventeenth  to  eighteenth 
segments ;  they  are  60  to  100  in  number;  cirrus-pouch  well  developed, 
may  attain  0.8  mm.  in  length, cirrus  spinous.  Female  organs:  Vagina 
ventral  of  cirrus-pouch;  ovary  visible  in  seventh  segment,  reaches  its 
highest  development  in  thirteenth  to  sixteenth  and  atrophies  in  twen- 
tieth to  twenty-second;  it  is  situated  slightly  to  the  right  of  the  median 
line;  receptaculum  seminis  globular;  the  transverse  uterus  appears  in 
fifth  or  sixth  segment,  develops  proximal  and  distal  pouches,  and 
begins  to  fill  about  the  fifteenth  segment;  eggs  oval,  88  /*  by  50  to  60  //. 


1  Specimens  distributed  from  collection  of  R.  Blanchard. 


154  TAPEWORMS  OF  HARES  AND  RAEJUTS— STILES.         VOL.XIX. 


Dorsal  canal  dorsal  to  lateral  of  ventral  canal.  Genital  canals  pass 
dorsally  of  longitudinal  canals  and  nerves. 

Host. — Horse  (Equus  caballus). 

This  diagnosis  is  based  in  part  upon  the  anatomical  discussion  of 
this  worm  by  Zschokke,1  in  1888. 

ANOPLOCEPHALA  TRANSVERSARIA  (Krabbe,  1879),  R.  Blanchard,   1891. 

(Plate  VI,  figs.  4-7.) 

1879,  Tcenia  transversaria  KRABBE,  Verb.  d.  K.  Ges.  d.  Frde.  d.  Natur,  Anthropol.  und 

Ethnographic,  XXXIV,  Moskau,  pp.  2-3,  figs.  1-6. 
1891,  AnoiilQcephala  transversaria  (KRABBE,  1879),  R.  BLANCHARD,  M6m.   Soc.  zool. 

France,  IV,  p.  448. 

Diagnosis. — Anoplocephala  transversaria  (Krabbe,  1879),  E.  Blanch- 
ard, 1891 :  Strobila  attains  10  to  10  cm.  in  length  by  G  to  8  mm.2  in 
breadth,  and  is  composed  of  200  to  300  segments.  Head  distinct,  0.6 
to  0.8  mm.  broad  by  0.6  mm.  long,  suckers  powerful  and  prominent; 
posterior  lobes  absent;  neck  absent.  Genital  pores  unilateral  at  about 
the  middle  of  the  margin'.  Male  organs :  60  to  80  testicles  to  each  seg- 
ment, confined  to  the  aporose  portion  of  the  median  field;  cirrus-pouch 
large.  Female  organs:  Yulva  ventral  of  cirrus-pouch;  receptaculum 
seminis  elongate;  ovary  in  pore-side  portion  of  median  field;  uterus 
transverse,  with  pouches.  Dorsal  canal  lateral  of  ventral  canal.  Gen- 
ital canals  pass  the  longitudinal  canals  and  nerves  dorsally.  Ova  with 
well-developed  pyriforni  body. 

Host. — Arctomys,  sp. ;  in  Turkestan  by  Fedschenko; 

Diagnosis  is  based  on  Zschokke's  anatomical  discussion  in  1888.3 

Genus  ANDRYA,  Railliet,  1893. 

1893,  Andrya,  RAILLIET,  Traitd  de  Zoologie  me'dicale  et  agricole,  I,  p.  283.     Type, 
Tcenia  rhopalocephala,  Riehm,  1881. 

Provisional  diagnosis. — Anoplocephaliua3  with  segments  broader  than 
long,  or  as  long  as  broad.  One  set  each  of  male  and  female  organs  to 
each  segment;  genital  pores  irregularly  alternate  with  marked  ten- 
dency to  unilateral ity;  female  glands  in  median  field  on  pore  side  of 
median  line;  uterus  appears  as  a  network  with  peripheral  dichotoinous 
branches  and  afterwards  assumes  a  more  saccular  form;  testicles 
confined  to  median  field;  a  distinct  round  or  elongate  pedunculated 
prostatic  gland  near  ventral  canal  on  pore  side  of  median  field;  genital 
canals  pass  dorsally  of  ventral  (and  ?  dorsal)  canal  and  nerve.  Dorsal 
canal  dorsal  to  dorso — ( 1  lateral )  of  ventral  canal.  Calcareous  corpus- 
cles develop  in  distal  portion  of  strobila.  Egg  with  well  developed 
pyriform  body,  the  horns  of  which  are  rather  short.  Hosts:  Eodents. 

Type. — Andrya  rhopalocephala  (Riehm,  1881),  Stiles,  1895. 

'Reches.  Strnc.  Anat.  et  Histl.  des  Cestodes,  Geneve,  pp.  18^6,  figs.  1-14. 

2There  is  evidently  a  typographical  error  in  Zschokke's  figures,  for  he  gives  the 
breadth  of  the  mature  segments  at  0.6  to  0.8  mm.,  yet  states  on  the  same  page  that 
the  proglottids  are  0.5  mm.  long,  and  twelve  times  as  broad  as  long. 

3  Reches.  Struc.  Anat.  et  Histl.  des  Cestodes,  Geneve,  1888,  pp.  47-63,  figs.  15-20. 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM. 


155 


The  genus  Andrya  was  proposed  by  Eailliet  in  the  following  words: 
II  n'est  pas  douteux,  bien  que  leur  e"tude  anatoiuique  soit  encore  peu  avanc6e,  que 
lea  anoploc6phalinos  des  Rongeurs,  pourvns  de  pores  genitaux  alternes,  doivent  otre 
se'pares  g<5n<5riquement  de  ceux  des  Equides,  qui  ont  les  pores  genitaux  imilate"raux. 
Nous  en  faisons  done  le  genre  Andrya,  d'apres  le  Tccnla  rhopalocephala  Riehm,  et  en 
1'honneur  de  Nicolas  Andry,  le  savant  me"decin  du  XVIIe  siecle,  qui  a  contribud  1'un 
des  premiers  a  dlucider  Thistoire  des  Teniades. 

Besides  the  type-species,  Eailliet  placed  here  Tcenia  rhopaliocephala^ 
Eiehm  (=  Anoplocephala  cuniculi,  E.  Blanchard)5  and  T.  wimerosa, 
Moniez.  The  latter  species  I  return  to  the  genus  Anoplocephala  (p.  151). 
In  my  preliminary  note  in  1895  I  placed  an  American  form  (Andrya 
americana)  in  Eailliet's  genus,  but  since  examining  Meyner's  specimens 
of  Bertia  mucronata  I  am  inclined  to  transfer  A.  americana  to  the  genus 
Bertia,  see  p.  165.  Eegarding  the  validity  of  the  genus  Andrya,  see  p. 
1G4.  Of  Eiehm's  original  material  I  have  obtained  the  following  speci- 
mens: 

Cotypes  of  Eiehm's  Tcenia  rhopalocephala  and  Tcenia  rhopaliocephala. 


U.  S.  N.  M. 
number. 

Label. 

Received  from  — 

Hedetermined  as  — 

Parasite. 

Host. 

1377 
1378 
1379 

1484 
1485 

Taenia  rhopalocephala. 
do             

Lepus  timidus  
Lepus  cuniculus.  . 

Vienna  Museum  .  . 
do  

Andrva  cuniculi. 
Do.? 
Andrya    rhopalo- 
cephala. 
Do. 
Do.? 

do 

do 

Taenia  rhopaliocephala 
Tsenia  rhopalocephala 

do  
do     

Leuctart 

do  

All  of  these  specimens  were  evidently  originally  determined  by 
Eiehm,  who,  however,  trusted  too  much  to  the  external  form  of  the 
segments,  for  a  careful  comparison  of  the  worms  with  Eiehm's  figures 
and  description  shows  that  some  of  the  specimens  were  misdetermined. 

Eiehm  states  that  his  T.  rhopalocephala  was  found  only  in  Lepus 
timidus^  while  T.  rhopaliocephala  (= Andrya  cuniculi}  was  confined  to 
Lepus  cuniculus.  Whether  the  discrepancies  between  his  labels  and 
this  statement  are  due  to  an  error  in  host  determination  or  an  error  in 
writing  the  labels  of  the  specimens  is  a  point  which,  of  course,  can  not 
now  be  settled.  For  the  present  I  adopt  his  statements  and  assume  an 
error  in  the  labels.  If  both  host  determination  and  labels  are  correct, 
then  Andrya  cuniculi  must  also  occur  in  Lepus  timidus.  This  point 
must  be  settled  by  new  collection  of  material. 

None  of  the  specimens  are  in  very  good  condition,  on  which  account 
I  am  unable  to  enter  into  a  detailed  study  of  the  organs.  My  observa- 
tions, however,  lead  me  to  accept  both  species  as  well  founded,  and  my 
results  agree  in  general  with  those  obtained  by  Eiehm. 

ANDRYA   RHOPALOCEPHALA   (Riehm,  1881),  Stiles,  1895. 

(Plates  VII,  figs.  1-7;  VIII,  1-3.) 
11800,  Alyselminthus pectinatus  (GoKZE,  1782),  ZEDER,  Erster  Nachtrag  z.  Naturg.  der 

Eingeweidewurmer,  Leipzig,  p.  246-249. 

?  1803,  Halysis  pectinata  (GoEZE,  1782),  ZEDER,  Anleitung  z.  Naturg.  der  Eingeweide- 
wiirmer,  p.  332. 


156  TAPEWORMS  OF  BARES  AND  RABBITS— STILES. 


1881,    Tcenia  rhopalocephala,  EIEHM,  Zeitschr.  f.  d.  ges.  Naturwiss.,  3  ser.,  VI,  p.  200; 

551-562,  pi.  v,  iigs.  1,  7-10,  18,  vi,  1. 
1891,  Anoplocepliala  rkopalocephala  (RiEHM,  1881),  R.  BLAXCIIARD.   Me"m.  Soc.  zool. 

France,  IV,  p.  448. 
1893,  Andrya  pectinata  ([GoEZE,  1782]  ZEDER.  1800),  RAILLIET,  Traite"  de  Zool.  m6d.  et 

agric.,  I,  p.  283. 
1895,  Andyra  rhopalocephctla  (RiEiiM,  1881),  STILES,  Vet.  Mag.,  II,  pp.  343,  344. 

I  can  find  no  evidence  that  Goeze  (1782)  had  any  single-pored  forms 
before  him  when  he  described  his  Tcenia  pectinata.  Zeder l  in  1800  rede- 
scribed  what  he  supposed  was  Goeze's  species,  but  states  that  the  pores 
were  single.  Eiehm2  in  1881  considered  Zeder's  species  identical  with 
the  one  now  under  consideration,  and  authors  have  followed  him  in 
this  opinion.  Zeder's  description  appears  to  me,  however,  altogether 
too  fragmentary  to  accept  this  view  as  proven ;  at  the  same  time  it  is 
impossible  to  definitely  disprove  Eiehm's  conclusion.  Eiehm3  in  1881 
was  the  first  to  recognize  Tcenia  rhopalocephala  as  a  distinct  form  $  his 
revised  diagnosis4  reads  as  follows : 

Kopf  hakenlos,  gross  und  keulenformig,  mit  stark  vorspringenden,  machtigen 
Saugnapfen,  scharf  abgesetzt  gegen  den  Halstheil.  GeschL •cktsoffnungen  einfach  im 
unteren  Viertheil  des  Proglottidenrandes  gelegen,  meist  durchgiingig  auf  derselben 
Seite.  Glieder  trapezforimg,  etwa  eben  so  breit  als  lang.  La'nge  im  ausgestreckten 
Zustande  60—80  cm.,  Breite  der  reifsten  Glieder  wenig  iiber  5  mm.  Wohnthier: 
Lepus  timidus. 

Blanchard5  in  1891  transferred  Eiehm's  species  to  the  genus  Anoplo- 
cephala,  while  Eailliet6  in  1893  took  it  as  type  of  the  genus  Andrya. 
Eailliet  reverted  to  Zeder's  specific  name,  but  I  now  adopt  rhopalo- 
cephala on  the  ground  that  Zeder's  pectinata  was  not  proposed  as  a  name 
for  a  new  species,  but  Zeder  was  under  the  impression  that  he  was 
redescribing  Goeze's  form.  Andrya  rlwpalocepliala  has  not  yet  been 
recorded  for  this  country.  The  following  statements  are  based  upon 
Eiehm's  specimen  (No.  1484,  U.S.N.M.),  mentioned  on  p.  155. 

The  anlage  of  the  genital  canals  and  female  glands  appears  earlier 
than  the  testicles.  In  the  lateral  third  of  the  segment  on  the  pore 
side  the  undifferentiated  anlage  of  the  canals  may  be  distinguished 
in  the  distal  portion  of  the  segment,  and  at  its  median  end  it  becomes 
widened  to  form  the  anlage  of  the  female  glands.  The  anlage  of  the 
canals  gradually  thickens  while  the  portion  destined  to  form  the  female 
glands  becomes  more  01*  less  distinctly  separated  from  it.  At  a  time 
when  this  apparent  separation  takes  place,  small,  quite  indistinct, 
points  of  chromatophile  material  appear  in  the  aporose  portion  of  the 
median  field  and  form  the  anlagen  of  the  testicles.  The  segment  now 

JErster  Nachtrag  zur  Natur.  der  Eingew.,  Leipzig,  pp.  246-249. 
2Stud.  an  Cestoden,  Zeit.  ges.  Nat.,  3  ser.,  VI,  pp.  545-546,  pi.  v-vi. 
3[Untersuchungen  an  den  Bandwiirmern  der  Hasen  und  Kaninchen],  Zeitschr.  ges. 
Naturw.,  3  ser.,  VI,  p.  200. 

4Studien  an  Cestodeo,  Zeitschr.  ges.  Naturw.,  3  ser.,  VI,  p.  551. 

6  Mem.  Soc.  zool.  France,  IV,  p.  448. 

6 Traite  de  Zool.  me"d.  et  agric.,  I,  p.  283. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM-  157 


measures  about  0.0  mm.  broad  by  0.35  mm.  long.  The  anlage  of  the  canals 
next  becomes  differentiated  into  the  anlagen  of  the  male  and  the  female 
ducts,  and  the  anlage  of  the  female  glands  divides  into  the  anlagen  of 
the  separate  glands.  The  testicles  are  now  distinct;  the  segments 
measure  1  mm.  broad  by  0.64  mm.  long.  The  ovarial  tubes  then  become 
distinct,  the  receptaculurn  seminis  appears  and  increases  in  size,  and  the 
elongated  prostata  and  the  cirrus  pouch  develop.  The  developed  cirrus 
pouch  measures  about  0.32  mm.  long  by  0.16  mm.  broad.  In  the  meantime 
the  testicles  have  encroached  upon  the  pore  side  of  the  segment  ante- 
rior to  the  ovary.  The  next  change  in  the  segments  is  brought  about 
by  the  development  of  the  uterus,  which  gradually  hides  the  testicles 
from  view.  As  iny  material  is  very  poorly  preserved,  and  hence  does 
not  stain  altogether  satisfactorily,  the  description  of  the  various  stages 
of  the  uterus  must  be  left  for  some  one  who  can  obtai>>  fresh  specimens. 
Plate  VII,  fig.  7,  represents  a  segment  measuring  2.08  mm. broad.  The 
cirrus  pouch  with  the  enclosed  inverted  cirrus  is  the  only  portion  of 
the  male  organs  which  is  now  visible.  The  female  glands  have  also  dis- 
appeared, but  the  receptaculum  seminis  is  still  present.  The  uterus 
fills  the  entire  median  field  of  the  segment  and  appears  as  a  reticulate 
structure,  with  dichotomous  branches  on  the  periphery.  As  the  ova 
develop,  the  boundaries  between  the  different  branches  of  the  network 
become  more  and  more  indistinct  and  the  uterus  assumes  the  form  of  a 
simple  sac  (Plate  -VIII,  fig.  1),  in  which  no  divisions,  or  at  most  extremely 
fragmentary  and  rudimentary  divisions,  are  visible.  The  cirrus  pouch 
and  the  receptaculum  seminis  are  still  present. 

Whether  this  uterus  is  primarily  an  actual  network,  or  whether  it  is 
originally  a  simple  tube  with  numerous  proximal  and  distal  branches 
which  secondarily  anastomose  and  then  gradually  disappear  to  form  a 
common  simple  sac,  can  not  be  stated  at  present  with  certainty,  but 
with  the  data  now  at  hand,  I  incline  decidedly  to  the  latter  view. 

The  ova  measure  52  to  60  /*;  the  pyriform  body  measures  12  ^  broad 
by  28  jj.  long  (horns  included). 

In  the  younger  segments  no  calcareous  corpuscles  are  visible.  When 
the  ova  in  the  uterus  become  more  distinct  and  the  partitions  in  the 
uterus  become  less  distinct,  a  few  calcareous  corpuscles  appear  in  the 
cortical  portion  of  the  segments.  As  the  shells  of  the  ova  develop,  the 
calcareous  corpuscles  become  numerous. 

From  Biehm's  anatomical  description  and  from  my  own  study,  I  pro- 
pose the  following  as  a  revised  specific  diagnosis : 

Diagnosis. — Andrya  rliopalocepJiala  (Eiehm,  1881),  Stiles,  1895:  Stro- 
bila  attains  60  to  80  cm.  in  length  by  5  mm.  in  breadth.  Head  unarmed, 
about  1  mm.  in  diameter,  nearly  quadrate  in  apex  view.  Neck  about 
1  mm.  long.  Segments  500  to  600  in  number;  active  segments  slightly 
broader  than  long;  segments  gradually  increase  in  length  so  that 
posterior  segments  are  as  long  as  broad;  they  may  attain  5  to  5.5  mm. 
in  breadth.  Genital  pores  single,  near  posterior  corner  of  the  segment, 


158  TA  PE  WORMS  OF  HARES  AND  RA  BBITS— STILES.         VOL.  xix. 

for  the  greater  part  unilateral.  Genital  organs  appear  in  about  the 
one-hundredth  segment.  Male  organs :  Testicles  dorsal,  comparatively 
few  in  number,  75  to  80  JJL  in  diameter,  and  more  numerous  in  the 
aporose  than  in  the  pore  side  of  the  segment 5  cirrus  pouch  0.32  to 
0.34  mm.  long  by  0.16  mm.  broad;  cirrus  short,  generally  lies  in  two 
spirals  within  the  pouch;  prostata  elongate.  Female  organs:  Vagina 
slightly  distal  of  cirrus  pouch,  swells  to. a  large  receptaculum  seminis 
median  and  dorsal  of  ventral  canal;  ovary,  shell-gland,  and  vitellogene 
gland  distal  in  pore  half  of  median  field:  uterus  ventral,  appears  as 
an  apparent  network  with  dichotomous  peripheral  branches,  and  finally 
forms  a  sac  with  indistinct  partitions  or  without  partitions.  Ova  52  to 
60  //;  pyriform  body  12  //  broad  by  28  /i  long.  Calcareous  bodies 
appear  in  segments  containing  ova,  and  become  numerous  in  segments 
in  which  the  egg  shells  are  distinct. 

Host. — European  Hare  (Lepus  timidus)  by  Biehm  in  Saxony. 

Cotypes.—Nos.  1379,  1484,  1485,  U.S.N.M.;  collection  of  Leupkart; 
Vienna  Museum. 

ANDRYA  CUNICULI  (R.  Blanchard,  1891),  Railli^t,  1893. 

Plates  VIII,  figs.  4-8;  IX,  fig.  1. 

1881,  Tceniarhopaliocephala  [nee  rhopalocephala]  RIEHM,  Zeitschr.  f.  d.  ges.  Naturwiss., 

3  ser.,  VI,  pp.  562-565,  pis.  v,  fig.  2,  vi,  fig.  3. 

1891,  Anoplocephala  cuniculi,  R.  BLANCHARD,  Me'm.  Soc.  zool.  France,  IV,  p.  447. 
1893,  Andrya  cuniculi  (R.  BLANCHARD,  1891),  RAILLIET,  Traite  d.  Zool.  me"d.  et  agric., 

I,  p.  283. 

Eiehm  described  this  form  in  1881  under  the  following  diagnosis: 

Kopf  hakenlos,  klein,  aber  gegen  den  sehr  diinnen  Halstheil  stark  keulenformig 
abgesetzt,  wenn  letztere  nicht  zu  stark  coutrahiert  ist.  Geschlechtsoffnimgen  ein- 
fach,  im  dritten  Viertheil  des  Proglottidenrandes  gelegen ;  Glieder  trapezformig, 
etwa  eben  so  lang  wie  breit.  Lange  im  ausgestreckten  Zustande  bis  100  cm.,  Breite 
der  reifsteu  Glieder  bis  zu  8  mm.  Wohnthier:  Lepus  cuniculus. 

Blanchard  in  1891  changed  the  name  to  cuniculi  on  grounds  of  arti- 
cles 54  to  55  of  the  International  Code,  and  placed  the  worm  in  the  genus 
Anoplocephala.  Eailliet  in  1893  placed  the  form  in  the  genus  Andrya. 

I  have  not  yet  found  this  species  in  the  United  States,  but  have  been 
fortunate  enough,  through  the  kindness  of  Geheimrath  Leuckart  and 
Dr.  von  Marenzeller,  to  obtain  some  of  Biehm's  original  stock. 

One  of  the  specimens,  which  was  mounted  whole,  shows  the  following 
details:  The  anterior  end  is  very  narrow  (0.4mm.)  and  segmentation 
is  scarcely  visible,  so  that  only  the  head  and  a  portion  of  the  neck  have 
been  lost.  Segmentation  is  noticed  0.64  mm.  from  the  anterior  extrem- 
ity, while  2  mm.  from  the  end  the  segments  are  perfectly  distinct,  meas- 
uring 0.8  mm.  broad  by  0.24  mm.  long.  The  anlagen  of  the  female 
organs  are  indistinctly  visible  at  about  this  point;  they  lie  close  to  the 
pore  side,  of  the  segment,  but  owing  to  the  poor  condition  of  the  mate- 
rial they  can  not  be  analyzed.  Testicles  could  not  be  distinguished  in 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  159 

these  segments,  but  they  appear  about  twenty  segments  later,  soon 
becoming  numerous  and  distinct  and  occupy  the  entire  median  field 
except  the  portion  taken  up  by  the  female  organs.  Plate  VIII,  fig.  6, 
represents  four  segments  about  15  mm.  from  the  anterior  end.  About 
forty  testicles  are  present.  The  anlage  of  the  female  glands  is  near  the 
median  line  in  the  pore  side  of  the  median  field;  it  is  roundish  and  not 
very  prominent.  The  genital  canals  are  not  very  distinct,  but  may  be 
traced  to  the  lateral  margin.  Plate  VIII,  fig.  7,  represents  three  seg- 
ments about  30  mm.  from  the  anterior  end,  The  testicles  have  increased 
in  number  and  size  ;  the  ovary  and  vitellarium  are  distinct,  but  the  shell- 
gland  is  scarcely  visible  j  the  genital  canals  have  become  well  differen- 
tiated, and  the  pore  has  developed.  The  greater  majority  of  the  genital 
pores  are  on  one  side  of  the  strobila,  and  are  situated  in  the  distal  half 
of  the  margin.  In  the  specimen  under  discussion  they  range  about  as 
follows : 

15.  16.  119.  44. 7.  5.  9. 

4.  4.  6.  6.  2.  2.  18.'etc* 

About  50  mm.  from  the  anterior  extremity  the  structure  of  the  seg- 
ment becomes  complicated  by  the  development  of  the  uterus.  My  prep- 
aration does  not  permit  a  study  of  its  gradual  development,  but  this 
organ  appears  to  be  much  more  complicated  than  one  would  expect 
from  Eiehni's  description.  At  first  no  distinct  limits  can  be  made  out, 
and  one  sees  only  numerous  ova  scattered  through  the  segment.  As  the 
testicles  atrophy,  however,  and  the  uterus  comes  more  distinctly  into 
view,  the  latter  is  apparently  composed  of  a  network  of  anastomosing 
tubes,  much  like  the  uteri  found  in  Moniezia,  arid  totally  different  from 
the  transverse  uterus  found  in  Cittotcenia,  Bertia,  and  Anoplocephala. 
As  in  the  case  of  the  anastomosed  testicles  of  Fasciola,  it  is  here  often 
impossible  to  distinguish  whether  we  are  dealing  with  anastomosing 
tubules  or  branched  tubules  which  lie  close  together. 

My  limited  material  will  not  warrant  a  more  minute  study  of  the 
uterus  of  this  form.  See  also  General  Eemarks,  p.  203. 

The  cirrus  pouch  is  seen  to  best  advantage  in  segments  in  which  the 
uterus  has  begun  to  develop.  It  is  pyriform  and  measures  0.4  mm.  long 
by  0.144  mm.  broad.  It  is  highly  muscular,  especially  in  its  proximal 
portion,  and  its  middle  portion  contains  a  prominent  dilatation  repre- 
senting a  vesicula  seminalis.  Median  to  the  pouch  is  found  a  roundish 
body,  which  evidently  corresponds  to  the  "prostata"  described  by 
Biehm. 

The  vagina  and  large  elongate  receptaculum  seminis  lie  distally  of 
the  pouch  and  vas  deferens,  as  Eiehm  has  already  described. 

The  ova  measure  48  to  60  //  in  diameter  j  the  pyriform  body  measures 
20  ^  broad  by  32  to  44  p.  long. 

The  following  is  proposed  as  a  revised  specific  diagnosis: 

Diagnosis. — Andrya  cuniculi  (K.  Blanchard,  1891),  Kailliet,  1893: 
Strobila  attains  100  cm.  in  length  by  8  mm.  in  breadth.  Head 


160  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 


unarmed,  about  0.5  min.  in  diameter;  rostellum  not  observed;  neck 
filiform.  Segments  500  to  800  in  number,  quadrate,  broader  than  long; 
gravid  segments  may  attain  8  mm.  in  breadth.  Genital  pores  in  about 
the  middle  of  the  margin  or  in  distal  half  of  the  margin,  irregularly 
alternate,  but  for  the  greater  part  unilateral.  Male  organs :  Testicles 
scattered  through  the  entire  breadth  of  the  median  field ;  cirrus  pouch 
0.4  to  0.48  mm.  long  by  0.14  to  0.16  mm.  broad,  quite  muscular,  and  con- 
tains a  distinct  vesicula  semiualis;  prostata  round.  Female  organs: 
Vagina  distal  of  cirrus  swelling  to  an  elongate  receptaculnm  seminis 
ventral  of  vas  deferens;  ovary  near  median  line  in  pore  side  of  median 
field:  uterus  forms  a  network  in  median  field,  but  as  the  ova  develop 
the  boundaries  of  the  meshes  become  quite  indistinct.  Ova  48  to  60  IJL 
in  diameter;  pyriform  body  20  //  broad  by  32  to  44  /.i  long.  Calcareous 
bodies  become  numerous  in  distal  segments. 

JTostf.— European  wild  rabbit  (Lcpus  cuniculus)  in  Saxony  by  Bielun ;  ? 
European  hare  (Lepus  timidus),  see  p.  155. 

Cotypes.~~Nos.  1377,  1378,  U.S.N.M.;  collection  of  Leuckart;  Vienna 
Museum. 

Genus  BERTIA,  R.  Blanchard,  1891. 

1891,  Bertia,  R.  BLANCHARD,  Me'm.  Soc.  zool.  France,  IV,  pp.  186-196.     Type,  Bertia 
studeri,  R.  Blanchard. 

Provisional  diagnosis. — Anoplocephaliuae,  with  segments  broader  than 
long.  Genital  pores  regularly  or  irregularly  alternate.  Uterus  (in  all 
cases  ?)  a  transverse  tube  with  proximal  and  distal  egg  pouches ;  geni- 
tal canals  pass  dorsally  of  dorsal  and  ventral  canal  and  lateral  nerve 
trunk,  but  in  the  two  cases  at  least  ventrally  of  dorsal  longitudinal 
nerve;  distinct  prostatic  gland  wanting.  Dorsal  canal  dorsal  to  dorso- 
lateral  of  ventral  canal.  Egg  with  well-developed  pyriform  body. 
Calcareous  corpuscles  present  or  absent.  Hosts:  Primates  and  rodents. 

Type.—B.  studeri,  K.  Blanchard,  1891. 

Two  years  prior  to  the  publication  of  the  genus  Andrya  by  Eailliet, 
R.  Blauchard,  in  1891,  proposed  the  genus  Bertia  for  anoplocephaline 
cestodes  with  alternate  genital  pores,  taking  B.  studeri  from  Antliro- 
popithecus  troglodytes  as  type  of  the  genus;  as  second  species  of  the 
genus  he  described  B.  satyri. 

Unfortunately,  on  account  of  paucity  of  material,  Blanchard  was 
unable  to  give  the  anatomy  of  the  type  species,  so  that  the  generic 
diagnosis  was  based  chiefly  upon  external  characters.  His  original 
diagnosis  reads  as  follows: 

Caput  crassum,  subsphaerieum,  rostro  aculeisque  carens,  acetabulis  ellipticis,  in 
dua  paria  valde  distautia  dispositis.  Collum  breve,  propre  tarn  larguni  quam  caput. 
Corpus  e  permultis  annulis  brevissiinis  latisque,  imbricatis,  constans.  Pori  geni- 
tales  raarginales,  tenuissimi,  ab  uno  annulo  ad  alterum  plus  minus  regulariter  alter- 
nantes.  In  anuulo  permaturo,  ova  in  plures  fasciculos  regulares,  transverse  dis- 
positus,  collecta.  Oncosphaera  pyriforme  apparatu  circumdata.  Involutio  ignota. 

From  Blanchard's  descriptions  of  the  species  the  following  may  be 
taken  as  provisional  specific  diagnoses. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  161 

BERTIA  STUDERI,   R.  Bianchard,   1891. 
(Plate  IX.,  figs.  2-3.) 

1891,  Bertia  studeri,  R.  BLANCHARD,  Me"m.  Soc.  zool.  France,  IV,  pp.  187-190,  figs.  1-4. 
1894,  Taenia  studeri  (R.  BLANCHARD,  1891),  BRAUN,  Vermes,  Bronn's  Klassen  und  Ord- 
nungen,  etc.,  IV,  36-37,  p.  1143. 

Diagnosis. — Bertia  studeri,  E.  Bianchard,  1891 :  Strobila  attains  130 
mm.  in  length,  15  mm.  in  breadth,  2.5  mm.  in  thickness;  contains  about 
400  segments.  Head  subspherical,  0.65  mm.  broad  by  0.61  mm.  long; 
suckers  oblong,  0.34  to  0.345  mm.  long  by  0.27  to  0.28  mm.  broad,  two 
arranged  on  dorsal  surface,  two  on  ventral  surface.  Neck  very  short 
(0.3mm.);  about  as  broad  as  head.  Segments  always  much  broader 
than  long;  maximum  breadth  15  mm.  at  45  mm.  from  head,  maximum 
length  about  0.35  mm.  Genital  pores  very  small,  lateral,  alternating 
very  regularly.  Male  organs :?  Female  organs :  Glands'?  Uterus  in 
fully  developed  stage  composed  of  30-35  polyhedral  packages,  0.5  to 
0.9  mm.  by  0.1  to  0.8  mm.,  arranged  in  transverse  row,  occupying  entire 
breadth  and  thickness  of  segments.  Ova  53  to  60  /*,  pyriform  body  14 
to  16  f.i  broad,  23  to  30  //  long,  horns  generally  straight,  oncosphere  10 
to  12  //.  Cortical  layer  of  strobila  supplied  with  numerous  calcareous 
corpuscles  15  to  20  yu  by  11  to  17  yu. 

Host. — Chimpanzee  (Anthropopithecus  troglodytes  (Linna3us)  [Troglo- 
dytes niger]))  by  Studer. 

Type. — Type  and  one  paratype  in  Zoological  Museum  at  Berne,  Swit- 
zerland. Fragments  in  collection  of  11.  Blauchard. 

BERTIA    SATYRI,  R.  Bianchard,  1891. 

1891,  Bertia  satyri,  R.  BLANCHARD,  Me~m.  Soc.  zool.,  France,  IV,  pp.  190-192. 
1894,  Tcenia,  satyri   (R.  BLANCIIARD,  1891),  BRAUN,  Vermes,    Bronn's   Klassen  und 
Ordnungen,  etc.,  IV,  36-37,  p.  1143. 

Diagnosis. — Bertia  satyri,  E.  Bianchard,  1891 :  Strobila  attains  245 
mm.  or  more  in  length  by  10  mm.  in  breadth  by  2  mm.  in  thickness,  and 
contains  about  350  segments.  Head  and  neck  unknown.  Segments 
always  much  broader  than  long,  attaining  a  maximum  length  of  0.75 
mm.  Genital  pores  very  small,  lateral,  irregularly  alternate.  Calcare- 
ous corpuscles  numerous,  attaining  30  by  20  yu.  Dorsal  canal  lateral 
of  ventral  canal.  Cirrus-pouch  claviform,  large  and  elongated.  Uterus 
resembles  somewhat  that  of  B.  studeri.  Ovum  35  to  38  /*  by  30  to  32  ^ 
pyriform  body  12  to  17  JJL  by  19  to  25  ^,  oncosphere  13  //. 

Host.— Oran-TJtan  (Simia  satyrus,  Linnaeus). 

Type. — Leyden  Museum. 

From  these  descriptions  it  is  impossible  to  come  to  any  satisfactory 
conclusion  as  to  whether  the  genera  Andrya  and  Bertia  should  be  kept 
separate  or  united.  The  form  of  the  segments  must  surely  be  rejected 
as  a  generic  character,  and  the  fact  that  the  pores  of  Bertia  show  a 
tendency  to  appear  regularly  alternate  while  those  of  Andrya  show 
Proc.  IN .  M.  vol.  xix— — 11 


162  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 

a  remarkable  tendency  to  unilaterality  can  not,  unassociated  with  other 
characters,  be  looked  upon  as  establishing  the  genera  as  distinct. 

In  the  preliminary  note  to  this  revision,  I  described  two  American 
parasites  as  provisional  members  of  the  genus  Andrya,  reserving  opin- 
ion as  to  the  validity  of  the  genus  and  calling  attention  to  some 
important  differences  in  the  American  and  German  leporine  single- 
pored  forms.  Since  the  publication  of  my  note,1  I  have  received  a  copy 
of  a  recent  paper  by  Meyuer  describing  two  new  species  of  cestodes, 
probably  allied  to  the  two  forms  of  Bertia  described  by  Blanchard,  and 
through  the  kindness  of  Geheimrath  Leuckart  I  have  obtained  one  of 
the  cotypes  of  Meyner's  Tcenia  (Bertia)  mucronata.  It  will  be  necessary 
to  consider  these  forms  briefly  in  connection  with  the  leporine  parasites. 

Meyner  evidently  accepts  Bertia  only  as  a  subgenus  of  Tcenia;  as 
Bertia  has,  however,  absolutely  no  generic  relations  with  Tcenia,  I  now 
change  his  specific  combinations  from  Tcenia  (Bertia)  mucronata  and 
T.  (B.)  conferta  to  Bertia  mucronata  and  B.  conferta. 

Meyner  discusses  the  anatomy  of  these  two  forms  in  detail,  and  from 
his  account  the  following  descriptions  may  be  taken  as  specific 
diagnoses : 

BERTIA  MUCRONATA  (Meyner,  1895),  Stiles,  1896. 
(Plate  IX,  figs.  4-5.) 

1895,   Tcenia  (Bertia)  mucronata,  MEYNER,   Zeitschr.  f.  Naturw.,  LXVIII,  (5  ser., 
VI),  pp.  1-86,  pi.  i,  figs.  1-7. 

Diagnosis. — Bertia  mucronata  (Meyner,  1895),  Stiles,  1896:  Strobila 
dagger-shaped,  attains  140  mm.  or  more  long  by  8  to  10  mm.  broad, 
serrate,  imbricate,  whitish  yellow.  Head,  0.34  to  0.714  mm.  broad, 
apex  nearly  square,  rostellum  wanting;  suckers  oval  0.255  mm.  broad, 
0.2  mm.  deep.  Neck  short,  not  sharply  separated  from  head.  Genital 
pores  irregularly  alternate.  Male  organs:  Testicles  appear  in  one 
hundred  and  twentieth  segment,  numerous,  75  to  100  p  in  'diameter, 
crowded  together  in  [antero  Jdorsal  portion  of  median  field.  Yas  defer- 
ens  dorso-anterior  of  vagina;  cirrus-pouch  not  mentioned.  Female 
organs:  Glands  in  pore  side  of  median  field;  vagina  long;  receptacu- 
lum  seminis  globular;  uterus  single,  transverse,  at  first  a  simple  tube 
appearing  in  about  one  hundred  and  thirtieth  segment;  eggs  enter  it 
in  three  hundred  and  fiftieth  segment  and  blind  pouches  are  formed. 
Ova  30  //,  with  three  membranes;  pyriform  body  15  to  16  yu  broad; 
oncosphere  13  to  14.4  //.  Three  longitudinal  nerves  each  side  of  seg- 
ment, of  which  middle  nerve  is  the  largest.  Dorsal  canal  dorsal  of 
ventral  canal.  Genital  canals  pass  dorsally  of  longitudinal  canals  and 
longitudinal  ventral  and  main  nerves,  but  ventrally  of  dorsal  nerve. 
Calcareous  bodies  4.9  to  18.4  //,  more  numerous  in  cortical  layer;  about 
150  visible  in  transverse  section  of  scolex,  10  to  12  visible  10  mm.  from 
anterior  extremity,  400  to  500  in  transverse  section  of  distal  segments. 

'Notes  on  Parasites— 38 :  Preliminary  note  to  "  A  Revision  of  the  Adult  Leporine 
Cestodes,"  Vet.  Mag.,  1895,  II,  p.  341-346. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  163 

Host. — Black  Howler  (Alouatta  caraya  (Humboldt)  [Mycetus  niyer]} 
in  Paraguay,  collected  by  Neumeister. 

Types. — In  collection  of  Leuckart;  one  cotype  No.  1483,  U.S.N.M. 

In  the  cotype  of  this  species  at  my  disposal  the  uterus  is  not  devel- 
oped. The  worm  presents  an  entirely  different  appearance  from 
Andrya.  The  muscular  cirrus  pouch  seems  to  be  almost  wanting,  so  far 
as  I  can  distinguish,  the  end  of  the  male  canal  appearing  simply  as  a 
widened  portion  of  the  vas  deferens  with  very  weak  muscles  and  much 
less  prominent  than  the  vagina.  The  prostatic  gland,  so  characteristic 
for  Andrya  rlwpalocephala  and  A.  cuniculi,  is  entirely  wanting.  Plate 
IX,  figs.  4-5,  give  the  general  topographical  anatomy  of  the  segment. 

I  do  not  believe  that  the  division  of  the  ovary  into  two  wings  as 
described  by  Meyner  can  be  maintained,  for  in  the  cotype  this  division 
is  extremely  irregular.  In  some  cases  the  ovary  is  not  divided ;  in  other 
segments  it  is  divided  into  two,  three,  or  four  wings.  I  further  find  the 
dorsal  canals  dorsal  to  dor  so-lateral  of  the  longitudinal  canals. 

BERTIA  CONFERTA  (Meyner,  1895),  Stiles,  1896. 

(Plate  IX,  fig.  6.) 

1895,  Tcsnia   (Bertia)  oonferta,  MEYNER,  Zeitschr.  f.  Naturw.,  LXVIII  (5  ser.,  VI), 
pp.  86-103,  pi.  i,  figs.  8-13. 

Diagnosis. — Bertia  conferta  (Meyner,  1895),  Stiles,  1896:  Strobila 
attains  84  mm.  or  more  in  length  by  6.5  mm.  in  breadth  ;  serrate.  Head 
roundish,  0.68  mm.;  rostelluin  wanting;  suckers  oval.  Neck  about  2 
mm.  long,  at  first  about  as  broad  as  the  head,  from  which  it  is  not 
sharply  separated.  Segments  always  much  broader  than  long;  sexual 
segments  measure  5.1  mm.  broad  by  0.27  mm.  long  by  1.02  mm.  thick. 
Genital  pores  irregularly  alternate.  Male  organs :  Testicles  numerous 
in  dorsal  portion  of  median  field  (in  figure,  of  transverse  section  con- 
fined to  aporose  two-thirds  of  median  field  at  plane  of  genital  pore), 
vas  deferens  dor  so -anterior  of  vagina,  its  lateral  portion  quite  wide; 
cirrus-pouch?  Female  organs:  Glands  in  middle  of  pore  side  of  seg- 
ment; receptaculum  seminisoval;  vagina  with  widened  ciliated  lateral 
portion.  Uterus  single,  transverse,  at  first  tubular,  later  with  blind 
sacs.  Ova  begin  to  enter  uterus  35  mm.  from  head  in  about  the  one 
hundred  and  thirtieth  segment.  Three  lateral  longitudinal  nerves  as 
in  B.  mucronata.  Dorsal  canal  dorsal  of  ventral  canal.  Genital  canals 
pass  dorsally  of  longitudinal  canals.  Calcareous  bodies  numerous,  may 
attain  19  /*;  20  to  25  visible  on  transverse  section  of  sexual  segments, 
600  to  800  on  transverse  section  of  distal  segments. 

Host. — Bonnet  Monkey  (Macacus  sinicus  (Linna3us)  [Macacus 
radiatus]). 

Type. — In  collection  of  Leuckart. 

While  looking  upon  Meyner's  paper  as  an  interesting  and  impor- 
tant contribution  to  the  knowledge  of  the  worms  of  this  group,  I  do 


164  TAPE  WOE  MS  OF  HAKES  AND  EABB1TS— STILES.         VOL.XIX. 

not  feel  justified  in  utilizing  liis  work  as  basis  for  more  than  a  provi- 
sional generic  diagnosis  for  Bertia  until  the  type  species  of  the  genus 
(B.  studeri)  can  be  studied  more  in  detail.  Nor  do  I  consider  the  data 
at  hand  sufficient  to  justify  a  worker  in  suppressing  Railliet's  genus 
Andrya  in  favor  of  the  earlier  genus  Bertia.  From  the  very  limited 
material  and  data  at  hand,  I  am  inclined  to  believe  that  Anoplocepkala*, 
Bertia ,  and  Andrya  will  all  eventually  be  recognized  as  good  genera, 
established  upon  well-recognized  anatomical  characters,  but  for  the 
present,  although  Anoplocephala  unquestionably  stands,  Bertia  and 
Andrya  can  be  accepted  only  as  provisional  genera,  and  as  convenient 
means  of  classification.  The  final  acceptance  of  the  genera  can  follow 
only  after  examination  of  a  larger  series  of  specimens  representing, 
if  possible,  more  species1  than  are  at  present  included  under  Andrya 
and  Bertia. 

To  utilize  the  generic  terms  Andrya  and  Bertia  provisionally  is  cer- 
tainly better  than  to  place  the  forms  in  the  same  genus  with  Tcenia 
solium. 

BERTIA  PLASTICA  (Sluiter,  1896),  Stiles,  1896. 

1896,  Tcenia  plastica,  Sluiter,  Centralbl.  f.  Bakteriol.,  Parasitenk.  u.  Infektionskr., 
I  Abt.,  XIX,  No.  24,  pp.  941-946,  figs.  1-6.     June  30,  1896. 

During  the  proof  reading  of  this  revision  an  article  has  appeared  by 
C.  Ph.  Sluiter,  describing  a  new  species  of  tapeworm  (Tcenia  plastica} 
from  Galeopithecus  volans.  Sluiter  is  inclined  to  consider  this  new  para 
.  site  as  very  closely  related  to  Anoplocephala  plicata,  A.  mamillana,  and 
A.  perfoliata,  all  of  which  he  retains  as  members  of  the  genus  Tcenia. 

Unfortunately,  several  important  points  in  the  topographical  anatomy 
have  been  omitted  by  the  author,  but  his  description  and  figures  show 
that  Tcenia  plastica  is  much  more  closely  related  to  Meyner's  Tcenia 
(Bertia)  mucronata  than  it  is  to  the  tapeworms  of  horses,  on  which 
account  I  transfer  it  to  the  genus  Bertia.  From  Sluiter's  description 
and  figures,  the  following  specific  diagnosis  is  written : 

Diagnosis. — Bertia  plastica  (Sluiter.  1896),  Stiles,  1896:  Strobilamore 


1  Gottheil  (1887)  described  two  other  cestodes,  which  may  have  some  bearing  upon 
the  question,  with  the  following  diagnoses : 

( Tcenia)  No.  1.  Length  20  cm.,  breadth  at  largest  segment  3.5  mm.  Head  globular, 
four  suckers,  no  rostellum,  and  no  hooks.  Neck  extremely  fine  and  filamentous — 2.5 
cm.  long.  Proglottides  slowly  increase  in  size,  greatest  breadth  being  only  attained 
15  cm.  from  head.  Sexual  orifices  at  the  sides.  Segments  oblong,  3.5  mm.  by  0.75 
mm.  From  Macaque  monkey  (Macacus  cynomolgm  (Schreber)  ). 

(Tcenia)  No.  2.  Length  of  largest  specimen,  15  cm.  Breadth  at  largest  segment, 
8  mm.  Head  large  and  clubbed,  four  suckers,  no  rostellum  or  hooks;  neck  short 
and  thick;  proglottides  rapidly  increase  in  size  after  the  first  inch  and  attain  their 
full  diameter  from  head,  7.5  cm.  mature  proglottides.  They  overlap  each  other  at 
their  posterior  angles.  Segments  mature  measure  8  by  4.5  mm. 

From  Macaque  monkey  (Macacus  cynomologus  (Schreber) )  and  Weeping  Capuchin 
(Cebus  capucinus  (Linnseus)  ). 

From  the  descriptions  and  figures  it  is  very  possible  that  these  two  forms  are 
anoploceph aline  cestodes,  but  Gottheil  gives  no  characters  which  will  aid  in  defi- 
nitely determining  the  question  at  hand. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  165 


or  less  lanceolate,  attaining  24  to  220  mm.  in  length  by  G  to  11  mm.  in 
breadth,  and  containing  80  to  400  segments.  Head  about  1  mm.  broad 
by  1  mm.  long;  suckers  round.  Neck  absent.  Proximal  segments 
increase  rapidly  in  breadth;  middle  segments  the  broadest;  distal 
segments  decrease  gradually  to  5  min.  in  breadth,  with  more  or  less 
crenate  and  imbricate  posterior  edge,  and  measure  0.5  to  0.76  mm.  long. 
Genital  pores  irregularly  alternate  in  about  the  middle  of  the  lateral 
margin;  genital  cloaca  and  organs  developed  in  fortieth  segment;  eggs 
in  uterus  in  fiftieth  segment.  Male  organs :  Cirrus  large;  cirrus-pouch 
muscular;  testicles  occupy  chiefly  the  anterior  portion  of  the  segments 
extending  the  entire  breadth  of  the  median  field.  Female  organs: 
Vagina  distal  to  cirrus-pouch;  receptaculum  seminis  elongate;  vagina 
and  receptaculum  together  extend  about  one-third  across  the  segment; 
ovary  very  broad,  extending  nearly  or  quite  to  the  aporose  submedian 
line;  shell-gland  and  vitollegene  gland  about  in  the  porose  submedian 
line;  uterus  tubular,  transverse  with  proximal  and  distal  pouches. 
Eggs  25  yw  in  diameter.  Topography  of  nerves  and  canals'? 

Host :  Flying  Lemur  ( Galeopitliecus  volans),  collected  by  Hubrecht  in 
In  India. 

Type:1*.  Deposited  in  Amsterdam.     Technique  of  type1? 

BERTIA  AMERICANA  (Stiles,  1895),  Stiles,  1896. 
(Plate  X,  figs.  1-10.) 

1855,  ?  Tcenia  laticephala,  LEIDY,  Proc.  Acad.  Nat.  Sci.  Phila.,  VII  (1854-1855),  Decem- 
ber, 1855,  p.  443. 
1895,  Andrya  americana,  STILES,  Vet.  Mag.,  Phila.,  II,  June,  p.  344.     Aug.  28, 1895. 

Leidy  has  given  a  short  description  of  a  tapeworm  (Tcenia  laticephala] 
from  the  Canada  Porcupine  which  agrees  in  some  characters  with  the 
form  I  described  (Andrya  americana)  from  the  yellow-haired  Porcupine. 
I  am  unable  to  find  Leidy's  types,  but  it  seems  to  me  very  questionable 
whether  the  two  parasites  are  identical.  Leidy's  description,  which  in 
a  measure  recalls  the  genus  Davainea,  reads  as  follows  : 

T^NIA  LATICEPHALA,  Leidy.  Head  large;  acetabula  opposite,  very  prominent, 
large,  hemispherical;  mouth  slightly  prominent,  unarmed.  Neck  short.  Anterior 
segments  of  the  body  short,  oblong  square;  posteriorly  square.  Generative  aper- 
tures marginal,  alternate.  Protruding  penes,  elongate  conical.  Length  of  one  speci- 
men 9  inches,  greatest  breadth  f  of  a  line.  Breadth  of  head  i  a  line ;  of  neck  £  a  line. 

Hob.  The  small  intestine  of  Hystrix  dorsata. 

B.  americana  was  described  in  the  preliminary  note  to  the  pres- 
ent revision  as  an  Andrya,  but  a  comparison  of  Meyner's  excellent 
anatomical  description  of  B.  mucronata,  and  of  his  cotype,  with  the 
form  under  discussion  shows  that  the  American  species  is  more 
closely  related  to  B.  mucronata  than  to  A.  rhopalocephala;  on  this 
account  A.  americana  is  transferred  in  the  present  paper  to  the  genus 
Bertia. 

Several  specimens  of  tapeworms  were  sent  to  the  collection  of  the 


166  TAPE  WOK  MS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 


Bureau  of  Animal  Industry  by  Professor  Elrod,  of  Bloomington,  Illi- 
nois, with  the  following  label :  "  From  Mesenteries  of  Canada  Porcupine. 
Snake  River,1  near  Nat.  Park,  Aug.,  1894,  M.  J.  Elrod." 

The  following  specific  description  will  easily  allow  the  recognition  of 
the  form : 

Diagnosis. — Bertia  americana  (Stiles,  1895),  Stiles,  1896:  Strobila 
attains  33  mm.  in  length  by  6  mm.  in  breadth  and  contains  about  90 
segments,  the  oldest  of  which  are  0.8  mm.  long.  In  some  specimens 
the  posterior  segments  become  much  narrower,  longer,  and  thicker  than 
the  middle  segments.  Head,  unarmed,  measures  0.6  mm.  broad  by  0.38 
mm.  long  by  0.32  mm.  thick,  and  is  nearly  rectangular  in  apex  view. 
The  neck  is  absent,  and  the  head  is  frequently  retracted  into  the  body, 
as  in  Drepanidotcenia  laceolata.  Suckers  round,  0.176  mm.  in  diameter, 
open  anteriorly.  Genital  anlagen  visible  in  the  earliest  segments. 
Genital  pores  alternate  in  posterior  half  of  margin.  Male  organs:  Tes- 
ticles form  a  continuous  band  in  the  distal  portion  of  the  median  field, 
extending  on  both  sides  to  the  longitudinal  canals;  about  seventy  testi- 
cles to  a  segment;  vas  deferens  runs  in  the  proximal  portion  of  the 
segment;  cirrus-pouch  lies  dorsal  of  the  vagina,  is  very  muscular,  0.48 
mm.  long  by  0.144  mm.  broad  and  extends  to  the  ventral  canal ;  it  con- 
tains a  vesicula  semiualis  (0.19  mm.  long)  in  its  proximal  portion,  and 
the  rather  short,  retracted,  spinous  cirrus  in  its  distal  portion.  Female 
organs:  The  anlagen  of  the  glands  are  seen  immediately  back  of  the 
head  in  or  near  the  median  line  ventral  of  the  testicles ;  at  first  the 
glands  are  but  little  differentiated,  but  on  their  pore  side  a  globular 
receptaculum  seminis  rapidly  develops  and  becomes  filled  with  sperma- 
tozoa; the  glands  develop  rapidly,  coming  to  lie  right  and  left  of  the 
median  line,  the  ovary  becoming  quite  broad.  The  development  of  the 
uterus  could  not  be  followed  in  detail,  but  eventually  it  occupies  the 
entire  median  field  and  becomes  filled  with  ova  40  n  in  diameter;  bulb 
of  pyriform  body  16  to  18//.  Excretory  and  nervous  systems:2  Dorsal 
canals  lie  lateral  of  ventral  canals  and  possess  a  thin  lining.  Genital 
canals  cross  the  longitudinal  canals  and  nerves  dorsally.  Calcareous 
corpuscles  absent. 

Hosts. — Yellow-haired  Porcupine  (Erethizon  epixanihus],  by  Elrod; 
Canada  Porcupine  (E.  dorsatus),  by  A.  K.  Fisher.3 

1  The  Canada  Porcupine  (Erethizon  dorsatm)  does  not  extend  so  far  west ;  the  host 
must  have  been  the  yellow-haired  Porcupine  (E.  epixanthus). 

2The  excretory  system  of  this  form  will  repay  a  careful  study;  in  several  anterior 
segments  (transverse  sections)  I  found  the  dorsal  canals  connected  with  the  trans- 
verse canals,  see  pi.  x,  fig.  8. 

•Since  finishing  this  paper  I  have  found  some  specimens  in  the  Bureau  of  Animal 
Industry  collection  (No.  1502)  which  agree  with  Elrod's  specimens,  and  bear  the 
label,  "E.  dorsatm."  These  specimens  were  collected  by  Dr.  A.  K.  Fisher  at  Lake 
George,  New  York.  Fisher  states  that  nearly  every  porcupine  he  has  examined  har- 
bors this  worm. 

Cobbold,  in  1862,  examined  some  parasites  from  the  same  host-species  and  deter- 
mined them  as  "Tcenia pectin ata,"  with  pores  "all  on  one  side." 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM. 


167 


Types. — Bureau  of  Animal  Industry  Cestode  Series  No.  1349  desig- 
nated as  type,  and  deposited  with  sections  of  paratypes  (Nos.  1350-1352) 
in  United  States  National  Museum.  Paratypes  distributed  as  follows: 
America:  Collection  of  Bureau  of  Animal  Industry;  collection  of  Stiles 
(U.S.N.M.);  collection  of  Hassall;  collection  of  M.  J.  Elrod  (Blooming 
ton,  Illinois).  Europe:  Berlin  Museum;  Vienna  Museum;  collection 
of  Bl  an  chard ;  British  (South  Kensington)  Museum  (London). 

Geographical  distribution. — North  America  (Wyoming,  by  Elrod,  and 
Lake  George,  New  York,  by  Fisher). 

BERTIA  AMERICANA   LEPORIS   (Stiles,  1895),  Stiles,  1896. 

(Plate  X,  figs.  11-15..) 
1895,  Andrya  americana  leporis,  STILES,  Vet.  Mag.,  II,  June,  p.  344.     Aug.  28, 1895. 

Of  this  form  I  have  but  five  specimens,  all  in  exceedingly  poor  con- 
dition and  unfit  for  any  histological  observations  or  detailed  anatomical 
study.  Enough  can  be  seen  on  the  preparations,  however,  to  give  a 
diagnosis  which  distinctly  separates  the  parasites  from  all  the  other 
forms  found  in  rabbits. 

The  specimens  were  collected  by  Cooper  Curtice;  when  they  came 
into  my  possession  there  was  nothing  upon  the  label  to  give  any  clue  to 
their  origin,  other  than  that  Curtice  collected  them  from  Lepus. 

The  specimens  (Bureau  of  Animal  Industry,  Cestode  Series  Nos. 
1170-1172,  1175-1176),  all  mounted,  measure  23  to  47  mm.  long,  the 
widest  segments  attaining  5  to  6.5  mm.  in  breadth.  Due  allowance  must 
here  be  made  for  the  contraction  of  the  specimens  and  the  fact  that 
they  were  subjected  to  pressure  in  mounting. 

The  head  is  present  on  all  specimens,  and  varies  in  measurement,  as 
shown  in  the  following  table : 

Measurements  of  heads  of  Bertia  americana  leporis. 


No.  (IT.  S.  N.  M.). 

Breadth. 

Length. 

1170      

mm. 
0.640 

mm. 

0.504 

1171                                                   

.656 

.448 

1172                                        

.656 

(?) 

1175                  

.688 

(?) 

1176  

.640 

.672 

The  form  of  the  head  can  not  be  taken  as  a  character  of  value,  as  it 
varies  according  to  contraction.  Plate  X,  figs.  12-13,  show  the  heads 
of  Nos.  1 170  and  1176,  U.  S.  N.  M.  Eostellum  could  not  be  distinguished. 
The  four  suckers  are  powerful,  and  open  diagonally  forward;  they  are 
unarmed  in  the  specimens  at  hand;  their  diameter  varies  between  0.22 
and  0.24  mm.,  the  muscular  wall  measuring  about  64  ^  thick.  There 
is  no  neck  present,  the  segments  being  perfectly  distinct  immediately 
back  of  the  head. 

The  number  of  segments  in  the  diiferent  specimens  varies  from  about 


168  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 

60  to  95,  but  in  no  case  was  the  strobila  complete,  as  all  of  them  had 
evidently  shed  proglottids.  In  some  cases  the  head  was  retracted  into 
the  anterior  segments,  as  is  described  for  Drepanidotcenia  lanceolata. 
In  only  one  specimen  (Plate  X,  fig.  13)  was  it  extended.  The  first 
segment  could  not  be  satisfactorily  measured;  the  second  segment 
(No.  1176,  U.S.N.M.)  measured  0.448  mm.  broad  (slightly  narrower  than 
the  head)  by  64  /*  long,  and  showed  no  trace  of  genital  organs.  The 
cirrus  was  distinctly  visible  in  one  specimen  in  the  sixth  segment,  but 
no  positive  statement  can  be  made  for  the  earlier  proglottids.  In  some 
of  the  other  specimens  the  cirrus  did  not  appear  until  several  segments 
later — in  the  eleventh  in  one  case.  The  poor  condition  of  the  material 
must,  however,  betaken  into  consideration  in  this  connection. 

Several  segments  after  the  appearance  of  the  cirrus,  a  roundish  body 
about  16  JJL  in  diameter  appears  in  each  proglottid,  alternating  a  very 
short  distance  to  the  right  and  left  of  the  median  line.  This  body, 
which  develops  first  from  the  eighth  to  the  fourteenth  segment,  evi- 
dently represents  the  receptaculum  seininis.  In  each  succeeding  seg- 
ment it  is  slightly  farther  from  the  median  line  than  in  the  segment 
immediately  preceding,  so  that  we  may  conclude  that  the  median  line 
of  the  cestrode  strobila  is  the  seat  of  very  active  growth,  a  conclusion 
supported  by  observations  on  other  species  also,  more  particularly  on 
the  early  genital  anlageu  of  Thysanosoma  giardi.1  The  testicles  soon 
appear  and  extend  in  an  irregular  line  across  the  median  field  of  the 
segment,  for  the  greater  part  distally  of  the  female  anlagen;  their  num- 
ber could  not  be  ascertained  exactly  because  of  the  poor  condition  of 
the  material,  but,  as  nearly  as  could  be  estimated  from  various  portions 
of  segments  which  were  in  better  condition,  there  are  about  50  testicles 
to  each  proglottid. 

The  genital  pores  are  alternate  and  situated  in  about  the  middle  of 
the  lateral  margin ;  the  genital  cloaca  is  generally  quite  deep.  The 
cirrus  and  cirrus  pouch  are  quite  characteristic  and  allow  an  immediate 
determination  of  the  form.  The  cirrus  pouch  assumes  various  shapes 
and  proportions  according  to  its  contraction,  but  in  general  may  be 
described  as  pyriform;  it  measures  about  0.4  mm.  long  by  0.16  mm. 
broad;  its  proximal  portion  (0.192  to  0.224  mm.)  is  extremely  muscular, 
being  provided  with  an  inner  circular  layer  and  an  outer  longitudinal 
layer,  the  two  together,  in  some  cases,  measuring  16  //  thick.  The  cirrus 
extrudes  from  the  pouch  in  the  majority  of  the  segments ;  when  retracted, 
it  appears  spinous  (wrinkles  of  the  cuticle?)  but  extruded  it  is  evidently 
smooth;  the  largest  cirrus  observed  was  0.24  mm.  long  by  32  /n  thick. 
The  continuation  of  the  cirrus  canal  in  the  proximal  portion  of  the  pouch 
is  swollen  into  a  vesicula  seminalis  and  from  the  proximal  extremity  of 
the  pouch  the  rather  prominent  vas  deferens  extends,  somewhat  sin- 
uously, through  the  anterior  portion  of  the  pore  side  of  the  segment  to 


1  A  Revision  of  the  adult  cestodes  of  Cattle,  Sheep,  and  allied  animals;  Bull.  4 
Bureau  of  Animal  Industry,  p.  63,  pi.  xn,  tig.  3. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  169 

the  vicinity  of  the  female  anl age— farther  than  this  it  could  not  be 
traced.  The  female  organs  gradually  undergo  a  change,  but  this  process 
can  not  be  described  in  detail  because  of  lack  of  proper  material.  All 
that  it  is  at  present  safe  to  say  is  that  a  dark  body  of  cells  arises  in  the 
vicinity  of  the  receptaculum  seminis  and  develops  into  the  female  glands 
very  similar  to  those  found  in  the  Bertia  of  the  porcupine;  the  uterus 
gradually  extends  over  the  segment  suppressing  the  genital  glands. 

Eggs  globular  with  three  membranes,  but  pyriform  body  not  visible; 
outer  membrane  40  to  42  u  in  diameter;  middle  membrane  could  not  be 
studied;  inner  membrane  immediately  surrounding  the  oncosphere  21 
JLI  in  diameter;  hooks  of  oncosphere  9  ^  long.  Numerous  small  calca- 
reous corpuscles  present  in  the  parenchyma. 

This  worm  has  an  entirely  different  appearance  from  the  one  found  in 
the  porcupine  and  it  is  possible  that  it  will  eventually  be  raised  to 
specific  rank.  My  material,  however,  does  not  warrant  such  a  step  at 
present. 

The  following  is  proposed  as  a  diagnosis  of  the  variety: 

Diagnosis.  —  Bertia  americana  leporis  (Stiles,  1895),  Stiles,  1896: 
Strobila  23  to  47  mm.  long  by  5  to  6.5  mm.  broad,  possessing  from  60 
to  95  segments.  Head  about  0.6  mm.  broad  by  0.4  to  0.6  mm.  long, 
unarmed;  suckers  large,  about  0.2  mm.  in  diameter,  unarmed.  Neck 
absent,  stobilization  beginning  immediately  back  of  the  head.  Genital 
pores  alternate,  situated  in  about  the  middle  of  the  lateral  margin. 
Genital  organs  develop  very  early,  the  male  organs  being  visible  some- 
times as  early  as  the  sixth  segment.  Cirrus  large  and  ( ? )  smooth. 
Cirrus  pouch  about  0.4  mm.  long  by  0.16  mm.  broad ;  its  proximal  por- 
tion is  very  muscular  and  contains  a  vesicula  seminalis.  Vas  deferens, 
large  and  prominent,  extends  from  the  region  of  the  female  glands  to  the 
cirrus  pouch  in  the  proximal  portion  of  the  segment ;  testicles  for  the 
greater  part  in  the  distal  portion  of  the  segment,  about  50  in  number, 
arranged  in  an  irregular  row  across  the  median  field.  Female  glands 
first  appear  in  the  eighth  to  fourteenth  segment  alternately  right  and 
left  of  the  median  line;  uterus  spreads  from  female  glands  and  occupies 
entire  median  field ;  ova  globular  with  three  thin  membranes;  pyriform 
body  apparently  not  present;  outer  membrane  40  to  42  }JL  in  diameter; 
middle  membrane?;  inner  membrane  21  /*  in  diameter;  hooks  of  onco- 
sphere 9  f.i  long.  Calcareous  corpuscles  present. 

Host. — Lepus,  sp.  ?,  by  Curtice,  development  unknown. 

Types. — Description  based  upon  five  specimens,  of  which  Bureau  of 
Animal  Industry  Cestode  Series  No.  1171  is  designated  as  Type  and 
deposited  in  the  United  States  National  Museum;  paratypes  Nos.  1170, 
1172,  1175,  1176  U.S.N.M. 

Original  material  poorly  preserved,  alcohol  ( ?)  method,  stained  with 
ha3inatoxylin. 

Geographical  distribution. — United  States  of  North  America  (State?) 
by  Curtice. 


1 70  TAPE  won  MS  OF  HARES  AND  RABBITS— STILES.      VOL.XIX. 


Genus  CITTOTCENIA,  Riehm,  1881. 

1881,  Cittotcenia,  EIEHM,  Zeitsclir.  f.  d.  ges.  Naturwiss.,  3  ser.,  VI,  p.  200.     Type, 

Cittotcenia  latissima  Riehm,  1881—  Tcenia  denticulata  Rudolphi,  1804. 
1881,  "Dipylidium,  LEUCKART,"  1863,  p.  p.  of  RIEHM,  Zeitschr.  f.  d.  ges.  Naturwiss., 

3  ser.,  VI.,  p.  200 ;  p.  565.     Type,  Tce.nia  canina  Liunreus,  1758. 
1891,  Moniezia  p.  p.,  R.  BLANCHARD,  Mdm.  Soc.  zool.  France,  IV,  p.  187;  p.  444. 

Type,  Tamia  expansa  Rudolphi,  1810. 
1893,  Ctcnotamia,  RAILLIET,  Traitd  Zool.  m6d.  et   agric.,  I,  p.  278.     Type,  Tamia 

marmotce,  Frolich,  1802. 

Diagnosis. — Cittotcenia,  Eiehm,  1881 :  Anoplocephalinre  with  segments 
broader  than  long  and  longer  than  thick ;  end  segments  in  some  cases 
showing  a  tendency  to  become  longer  and  narrower.  Two  full  sets  of 
genital  glands  and  two  lateral  genital  pores  to  each  segment;  gener- 
ally one,  in  some  cases  two,  simple  transverse  tubular  uteri  in  each 
segment;  uterus  generally  possesses  simple  proximal  and  distal  diver- 
ticula.  The  vagina  is  ventral  of  the  cirrus  pouch  on  both  sides  of  the 
segment.  Dorsal  canal  varies  somewhat  in  position,  but  shows  a  con- 
stant tendency  to  lie  between  the  ventral  canal  and  nerve,  especially 
at  the  plane  of  the  genital  pores.  Genital  canals  cross  the  longitudinal 
canals  and  nerves  dorsally.  Interproglottidal  glands  absent.  Calca- 
reous bodies  not  yet  recorded.  Eggs  with  well-developed  pyriform 
body,  the  horns  of  which  are  long,  generally  filamentous,  crossing  each 
other. 

Hosts.— Eodents. 

Type. — Gittotamia  latissima,  Eiehm,  1881,  =  Cittotcenia  denticulata 
(Eudolphi,  1804),  Stiles  &  Hassall,  1896. 

Eiehm  proposed  the  genus  Cittotcenia  in  1881,  with  C.  latissima  as 
type  and  only  species ;  his  description  reads  as  follows : 

4.  Cittotcenia  latissima.  Die  Gattung  ist  errichtet  mit  Riicksicht  auf  die  merkwiir- 
dige  Ausbildung  des  Excretionsorganes,  welches  nicht  leiterartig,  sondern  mit  je 
3-5  vielfach  omastomosirenden  und  in  einander  iibergehenden  Hauptseitenstammen 
und  zahlreichen,  das  ganze  Thier  netz-formig  durchsetzenden  Nebeniisten,  ranken- 
artig  die  Tame  dnrchzieht.  C.  latissima  hat  doppelte  Geschleehtsorgane,  welche 
auf  den  zizzenartig  vorspringenden  Hinterecken  der  Glieder  nach  aussen  miinden. 
Der  Kopf  breiter  als  bei  den  vorigen  erscheint  vorn  abgestutzt,  hakenlos.  Liinge  der 
Strobila  bis  2£  Fuss,  Breite  der  letzten  Proglothiden  bis  iiber  f  Zoll.  Wohnthier: 
Kaninchen. 

In  the  same  paper  Eiehm  placed  two  other  double- pored  anoploce- 
phaline  cestodes  in  Leuckart's  genus  Dipylidium,  i.  e.,  D.  pectinatum  and 
D.  leuckarti,  and  later  in  the  same  year l  he  rejected  his  newly  estab- 
lished genus  Cittotcenia,  uniting  it  with  Dipylidium. 

In  1891  Blanchard 2  united  in  the  genus  Moniezia  the  double-pored 
anoploceph aline  cestodes  of  rodents  with  those  of  ruminants.  In  1893 

1  Studien  an  Cestoden.     Zeitschr.  f.  d.  ges.  Naturwiss.,  3  ser.,  VI,  1881,  pp.  565,  580. 

2  Mem.  Soc.  zool.  France,  IV,  1891,  p.  187. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MI'S  BUM.  171 


Stiles '  excluded  the  known  parasites  of  rodents  from  the  genus  Moniezia, 
while  Railliet  proposed  for  the  double-pored  leporine  forms  the  genus 
Ctenotccnia  with  Tcenia  marmotw  as  type. 

At  first  thought  this  genus  does  not  appeal  to  helminthologists,  who 
have  not  paid  special  attention  to  the  Anoplocephalin*,  as  being  well 
founded,  but  an  anatomical  study  of  the  various  forms  does,  I  believe, 
fully  justify  the  separation  of  the  double  pored  cestodes  of  rodents 
from  the  double  pored  forms  found  in  ruminants.  The  generic  term 
Cittotcenia  having  priority  must  of  course  be  accepted  in  preference  to 
Ctenotamia. 

The  characters  of  the  three  genera  of  double-pored  anoplocephaline 
forms,  according  to  my  present  interpretation,  are  as  follows: 

Diagnosis. — Moniezia,  R.  Blanchard,  1891:  Anoplocephaline  with 
segments  generally  broader  than  long  and  longer  than  thick ;  end  seg- 
ments showing  a  tendency  to  become  longer  and  narrower.  Two  full 
sets  of  genital  organs,  with  two  (very  complex)  reticulate  uteri  and  two 
lateral  pores  in  each  segment.  On  the  right  side  the  vagina  is  ventral, 
cirrus  dorsal;  on  the  left  side  vagina  dorsal,  cirrus  ventral.  Dorsal 
canal  lies  dorso-inedian  of  ventral  canal.  Genital  canals  cross  the  lon- 
gitudinal canals  and  nerves  dorsally.  Interproglottidal  glands  gener- 
ally present.  Calcareous  corpuscles  absent  from  parenchyma.  Eggs 
with  well  developed  pyriform  body,  the  horns  of  which  generally 
(always  ?)  end  in  a  disk. 

Hosts. — Ruminants. 

Type. — Moniezia  expansa  (Eudolphi,  1810),  E.  Blanchard,  1891. 

For  diagnosis  of  Cittotcenia,  see  above,  p.  170. 

Diagnosis. — Thysanosoma,  Diesing,  1835:  Auoploeephalma3  with  seg- 
ments generally  broader  than  long  and  longer  than  thick;  end  seg- 
ments show  a  tendency  to  become  longer  and  narrower.  Two  sets  or 
one  set  of  genital  glands  and  two  lateral  genital  pores,  or  one  lateral 
genital  pore  to  each  segment;  one  transverse  undulating  uterus  with 
ascon-spore  or  cornucopia-like  egg-pouches.  Genital  canals  cross  the 
ventral  canal  and  nerve  dorsally,  the  dorsal  canal  ventrally.  Interpro- 
glottidal glands  absent.  Calcareous  bodies  absent  from  the  paren- 
chyma. Horns  of  pyriform  body  absent. 

Hosts. — Rum  inauts. 

Type. — Thysanosoma  actinioides,  Diesing,  1835. 

From  this  analysis  of  characters  it  will  be  seen  that  Cittotcenia  forms 
an  excellent  intermediate  genus  between  Moniezia  and  Thysanosoma. 

The  genus  may  be  divided  into  two  groups :  Marmotce  or  Denticulata 
Group  and  Pectinata  Group,  the  division  being  based  upon  the  form  of 
the  cirrus  pouch.  In  the  first  group  this  organ  is  pyriform,  distinct  and 
very  muscular ;  in  the  second  group  it  is  more  elongated,  resembling  the 
nozzle  to  a  hose,  and  is  less  distinct. 

'Centralbl.  f.  Bakteriol.  u.  Parasitenk.,  XIII,  1893,  p.  457;  also,  Bull.  4,  Bureau 
Animal  Industry,  1893,  p.  54. 


172  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 


MARMOTS  OR  DENTICULATA  GROUP. 

CITTOT^ENIA   MARMOTS  (Frolich,  1802)  Stiles  &  Hassall,  1896. 
(Plate  XI,  figs.  1-8 ;  Plate  XII,  figs.  1-2. ) 

1802,  Tcenia  marmota,  FROLICH,  Der  Naturforscher,  XXIX,  p.  77-79,  pi.  n,  fig.  17-20. 
1891,  Moniezia  marmota;  (FROLICH,  1802),  R.  BLANCHARD,  M6m.  Soc.  zool.  France,  IV, 

p.  187 ;  pp.  444, 461-467,  tigs.  33-35. 
1893,  Ctenotwnia  marmota}  (FROLICH,  1802),  RAILLIET,  Trait6  d.  Zool.  m6d.  et  agric., 

I,  p.  278. 
1896,  Cittotcvnia  marmota  (FROLICH,  1802)  STILES  &  HASSALL,  Veterinary  Magazine, 

III,  p.  407. 

Gceze l  in  1782  examined  a  worm  which  Blunienbach  had  collected 
from  the  Marmot;  this  parasite,  which  was  probably  Gittotamia  mar- 
motce,  Goeze  considered  identical  with  his  Tcenia  pectinata. 

Frolich2  in  1802  found  a  cestode  in  the  intestine  of  the  Marmot  (Arc- 
tomys  marmota)  and  described  it  under  the  name  Tcenia  marmotce,  as 
follows : 

[p.  77.]  43 :  Taenia  Marmoiae  oblonga  lanceolata  antice  attenuata  utrimque  serrata, 
articulis  brevissimis,  medio  utrimque  poro  pilifero  notatis,  capite  mutico.  Tab.  II, 
f.  17-20. 

MURMELTHIER-BANDWURM. 

Die  Lange  dieser  "Wiirmer  ist  sehr  veranderlich.  Die  grossten  Individuen  batten 
6-7  Zoll  in  der  Lange,  ungefabr  II'"  in  der  Breite  am  Hintertheile,  und  beinabe  1'"  in 
der  Dicke.  Ibre  Gestalt  ist  iiberbaupt  lanzettformisr,  in  die  Liinge  gezogen,  am 
Vordertbeile  allmalig,  aber  sebr  betriicbtlicb,  verscbmiilert,  so  dass  das  Vorderende 
am  Gruude  des  Raises  kaum  etwas  iiber  V"  breit  ist.  Aeltere  Wiirmer,  die  durcb 
Geburten  scbon  mebrere  Glieder  abgesetzthaben,  sind  am  Hinterende  am  breitesten, 
abgestutzt,  oder  balbmondformig  ausgerandet;  jiiugere  Wiirmer,  die  nocb  nicht 
gebobren  haben,  werden  am  Hinterende  etwas  schmacbtiger,  uud  das  letzte  Glied 
ist  abgerundet. 

Der  Kopf  ist  verhultnissmlissig  sebr  klein,  rundlicb,  riissellos,  unbewaffnet,  bell- 
weiss,  mit  vier  deutlicben  Saugblasen,  die  gepaart  iiber  einanderstebeu;  [p.  78]  an 
der  Spitze  ein  undeutlicbes  Knotchen,  statt  des  Riissels. 

Der  Hals  ist  zusammengedriickt  rundlicb,  sebr  scbmiicbtig,  etwas  lunger  als  der 
Kopf,  gliederlos,  abwiirts  breiter,  und  unvermerkt  in  den  Vorderkorper  iibergebend. 

Das  Vorderende  des  Korpers  scheint  da,  wo  es  an  den  Hals  grenzet,  fast  gliederlos, 
wird  von  da  abwtirts  sanft  uud  unvermerkt  breiter,  und  die  Glieder,  die  vorwiirts  nur 
durcb  Querstreifen  angezeigt  waren,  entwickeln  sicb  da  deutlicher. 

Die  Glieder  des  Korpers  sind  ganz  sicbtbar  ineiuander  geschoben,  wenigstens  20 
mal  breiter  als  lang,  gleicbbreit,  dicklicb,  perlenweiss,  zu  beiden  Seiten  in  eineu 
scharfen,  bervorragenden  Zabu  auslanfend,  in  der  Mitte  der  Oberfliiche  eine  einge- 
clriickte  Langslinie.  Jedes  Glied  bat  in  der  Mitte  des  Seitenrandes  ein  duukleres, 
hervorragendes,  wulstiges,  in  der  Mitte  durcbbobrtes  Knotcben,  aus  welcbem  ein 
baarfeiner,  geradegestreckter,  scbneeweisser,  iiber  V"  langer  Borsteii  bervorragt. 
Diese  Seitenborsten  sind  nicbt  iinmer  sichtbar,  sondern  sebr  oft  in  den  Rand  der 
Glieder  zuriickgezogen,  wo  sie  denn  an  jeder  Seite  des  Gliedes  eiue  bellere  Querlinie 
bilden.  Die  bintern  Glieder  nebmen  an  Breite  und  Liinge  etwas  zu. 

1  Versuch  einer  Naturg.  der  Eiiigeweidewiirmer  tbieriscber  Korper,  Blankenburg, 
p.  363,  footnote. 
3  Beitrage  zur  Nat.  der  Eingeweidew.,  Der  Naturforscbjer,  XXIX,  1802,  Halle,  p.  77. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  173 

Wolmort:  In  den  diinnen  Gedurmen  des  Murmeltbiers  (Arctomys  marmota  Scbreb.). 

Anm.  Icli  traf  diese  scbone  Wurmart  in  dem  genannten  Thiere,  das  in  den  all- 
gauischeu  Alpen  einbeimiscb  ist,  und  von  dem  icb  ein  paar  Stiicke  am  28  Julius  1797 
zu  untersucben  Gelegenbeit  bekam,  in  ausserordentlicber  Menge  an.  In  den  Gedilrm- 
en  ernes  iiltern  Murmeltbiers  fand  icb  ganze  Nester  von  diesen  Wurmern,  so  dass 
ibrc  Zabl,  im  Ganzeu  genommen,  iiber  50  Stiicke  betragen  haben  muss.  Es  lagen 
ibrer  niancbmal  5-7  beysammen,  und  debnten  den  Darnikanal,  indem  sie  in  Knaulen 
iibereiuander  lagen,  ungemein  aus. 

Icb  sab  Stiicke  von  diesen  Wiirmern,  deren  bintere  Glieder  in  der  Mitte  durch- 
locbert  waren :  andere  Individuen  batten  sich  am  Hintertbeile  durcb  die  Scblin<»-e 

7  O 

gescboben. 

Man  siebt  von  selbst,  dass  die  gegenwartige  Art  mit  dem  lanzetfdrmigen  Baud- 
wurme1  ungemein  viele  Aebnlicbkeit  babe,  da  aucb  dieser  an  den  Randern  der  Glie- 
der kurze  Seitenborsten  zu  naben  pflegt ;  demungeacbtet  kanu  icb  mich  bey  genauer 
Vergleichung  uicbt  iiberzeugen,  dass  beide  Wiirmer  eiuerley  Art  seyn.  Aucb  die 
Tcenia  pcctinata-  ist  mit  dieser  Art  nabe  verwandt. 

Eudolplii3  in  1814  and  1819,  Diesing4  in  1850,  and  Baird5in  1853, 
mention  Frolich's  parasite  under  Tcenia  pectinata,  accepting  Tcenia  mar- 
motcu  as  synonym.  E.  Blanchard 6  in  1891  was  evidently  the  second 
zoologist  to  find  this  species;  he  found  214  worms  in  four  marmots  at 
Briancon,  in  September,  1887.  The  specimens  attained  112  mm.  in 
length  by  5  to  13  mm.  in  breadth.  Head  measured  0.55  to  0.63  mm. 
long  by  0.80  to  0.84  mm.  broad ;  suckers  globular,  120  ^  in  diameter; 
neck  absent;  segments  varied  from  56  ^  to  0.75  mm.  in  length;  genital 
pores  double  and  opposite,  in  posterior  half  of  lateral  margin.  Blanch- 
ard  described  a  subcuticular  excretory  system,  with  two  lateral  longi- 
tudinal canals,  connected  by  a  transverse  canal  with  the  canals  of  the 
opposite  side.  Ova  measured  48  to  60  yu;  bulb  of  pyriform  body,  23  /*. 

Stiles7  in  1893  found  that  the  dorsal  canal  lies  between  the  ventral 
canal  and  nerve,  and  that  the  genital  canals  run  dorsally  of  the  longi- 
tudinal canals  and  nerves.  He  was  unable  to  find  Blanchard's  acces- 
sory excretory  system.  Later  he8  figured  the  genital  organs. 

Railliet9  in  1893  took  this  species  as  type  of  his  new  genus  Cteno- 
tcenia. 

From  writings  of  Frolich  and  Blanchard  and  from  my  own  studies 
I  propose  the  following  as  revised  specific  diagnosis: 

Diagnosis. — Cittotcenia  marmotw  (Frolich,  1802),  Stiles  &  Hassall, 
1896:  Strobila  attains  112  mm.  (perhaps  more)  in  length  by  5  to  13  mm. 
in  breadth.  Head,  0.8  mm.  broad  by  0.5  to  0.6  mm.  long ;  hooks  absent ; 


1  Goeze,  Eingeweidew.,  p.  377,  pi.  xxix,  fig.  3-12. 
3Ebeudas,  p.  363,  pi.  xxvn,  fig.  7-12. 

3Erster  Nacbtrag,  etc.,  Ges.  Naturf.  Freunde  Berlin,  etc.,  VI,  1814,  p.  108;  Ento- 
zoorum  Synopsis,  1819,  p.  488. 
4Systema  belmintbum,  I,  p.  498. 

6  Cat.  Specs.  Entozoa  or  Intestinal  Worms,  Coll.  Brit.  Mus.,  p.  78. 
6  Mem.  Soc.  zool.  France,  IV,  pp.  444,  461-466,  figs.  31-35. 
7 Cent.  f.  Bakt.  u.  Paras.,  XIII,  pp. '449-450,  fig.  9. 

8  Stiles  &  Hassall,  1893,  Bull.  4,  Bureau  of  Animal  Industry,  p.  71,  pi.  vn,  figs.  6-7. 
9Traite  de  Zool.  me'd.  et  agric.,  I,  p.  278. 


174  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.         VOL.XIX. 

rostellum  not  visible.  Keck  absent,  segmentation  beginning  immedi- 
ately back  of  the  head,  the  proglottids  rapidly  becoming  distinct; 
segments  vary  in  length  from  56  /t  to  0.75  mm.,  and  are  always  much 
broader  than  long,  measuring  in  some  cases  13  mm.  broad;  the  poste- 
rior flap  projects  but  a  short  distance  over  the  anterior  border  of  the 
next  following  segment.  Genital  pores  double  and  opposite  in  poste 
rior  half  of  lateral  margin.  Genital  cloaca  quite  deep.  Anlagen  of 
genital  canals  and  female  glands  visible  in  the  first  segments,  testicles 
appear  a  little  later.  Male  organs:  Testicles  appear  about  the  thir- 
tieth segment,  between  100  and  150  in  number  to  a  proglottid,  scat- 
tered through  the  median  field  between  the  ovaries;  cirrus-pouch,  0.5 
mm.  long  by  0.17  mm.  broad,  very  muscular,  with  vesicula  seminalis  in 
proximal  portion  and  coiled  smooth  penis  in  distal  portion.  Female 
organs:  Female  glands  nearly  1  mm.  from  the  lateral  border  of  the 
segment,  some  distance  from  the  longitudinal  canals;  ovarian  tubules 
appear  about  the  thirty-fifth  segment,  reach  their  highest  development 
from  the  forty-seventh  to  fifty-seventh  segments  and  then  rapidly  atro- 
phy, disappearing  almost  entirely  by  the  sixtieth  segment;  vagina 
ventral  of  the  cirrus-pouch  on  both  sides  of  the  segment;  uterus  single, 
transverse,  proximal  to  testicles,  and  possesses  proximal  and  distal 
blind  pouches,  similar  to  those  of  Cittotcenia  pectinata.  Ova,  48  to  00  //  ; 
bulb  of  pyriform  body,  23  //.  Excretory  system :  Dorsal  canal  between 
ventral  canal  and  nerve.  Longitudinal  nerves  rather  close  to  lateral 
margin,  near  distal  end  of  cirrus-pouch.  Genital  canals  cross  the 
longitudinal  canals  and  nerve  dorsally. 

Host. — Marmot  (Arctomys  marmota)  by  Frolich  and  Blanchard. 

Types. — Original  type  (?).  Typical  specimens  with  R.  Blanchard 
(Paris),  collection  Bureau  of  Animal  Industry  (No.  1370,  B.A.I.),  and 
collection  of  Stiles  (U.S.N.M.). 

Geographical  distribution. — (!)  Frolich;  France  (Brian§on),  by  Blan- 
chard. 

CITTOT^ENIA  DENTICULATA  (Rudolphi,  1804),  Stiles  &  Hassall,  1896. 
(Plate  XII,  figs.  3-8;  Plate  XIII,  figs.  1-3.) 

1804,  Tcenia  denticulata,  RUDOLPHI,  Bemerkungen  a.  d.  Gebiete  d.  Naturg.,  etc.,  p.  81. 

1828,  Alyselminthus  denticulatus  (RUDOLPHI,  1802),  DE  BLAINVILLE,  Diet.  d.  Sci.  nat., 
LVII,  p.  607. 

1853,  Tcenia  goezei,  BAIRD,  Cat.  Entozoa  Coll.  British  Museum,  p.  78. 

1881,  Cittotcenia  latissima,  Reihm,  Zeitschr,  f.  d.  ges.  Naturw.,  3  ser.,  VI,  p.  200. 

1881,  Dipylidium  latissimum  (Riehm,  1881),  RIEHM,  Zeitschr.  f.  d.  ges.  Naturwiss.,  3  ser., 
VI,  pp.  583-590,  pi.  v,  figs.  5, 15, 17;  vi,  fig.  2. 

1887,  Tcenia  latissima  (RIEHM,  1881),  NEUMANN,  Trait6  des  maladies  parasitaires  non- 
mi  crobiennes,  p. 426. 

1891,  Moniezia  denticula  \ta]  (RUDOLPHI,  1804),  R.  BLANCHARD,  Me"m.  Soc.  zool.  France, 
IV,  p.  187. 

1891,  Moniezia  latissima  (RIEHM,  1881),  R.  BLANCHARD,  Me"m.  Soc.  zool.  France,  IV, 
pp.  187, 451. 

1891,  Moniezia  goezei  (BAIRD,  1853),  R.  BLANCHARD,  M6m.  Soc.  zool.  France,  IV,  pp. 
444, 452-457,  figs.  21-25. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  175 

1893,  Ctenotwnia  goezei  (BAIKD,  1853),  RAILLIET,  Trait<5  d.  Zool.  mc~d.  et  agric.,  T,  p.  278. 
1896;  Ctenotcenia  denticulata  (RUDOLPHI,  1804),  STILES  &  HASSALL,  Vet.  Mag.,  Ill,  p. 

6-9,  and  Centralbl.  f.  Bakt.,  Parasitenk.  u.  Irifektionskrankh.,  Erste  Abt., 

XIX,  70-72. 
1896,  Cittotcenia  denticulata  (Rudolphi,  1804),  Stiles  &  Hassall,  Vet.  Mag.,  Ill,  p.  407. 

This  species  has  had  a  most  curious  history,  which  should  be  a  demon- 
stration to  every  helminthologist  not  yet  convinced  that  in  determining 
a  cestode  the  microscope — not  the  yardstick — is  his  most  important 
instrument,  and  that  internal  topographical  anatomy — not  conditions 
of  contraction  of  the  external  form — furnishes  the  important  characters 
for  classification. 

At  present  we  may  be  said  to  have  two  extreme  parties  in  helmin- 
thological  work;  one  party  apparently  considers  minute  histological 
details  as  all-important,  and  looks  down  upon  the  systematic  work, 
especially  that  class  of  systematic  work  which  studies  into  the  biblio- 
graphic history  of  the  species  and  demands  a  consistent  application  of 
the  international  rules  of  nomenclature;  a  second  party  appears  to 
conceive  the  highest  helminthology  as  consisting  of  determining  and 
describing  species  chiefly  upon  external  form,  looking  upon  histological 
and  anatomical  details  as  "of  interest  to  those  who  are  interested  in 
such  matters,"  but  of  no  importance  to  science.  The  history  of  C.  denti- 
culata should,  however,  convince  everyone  that  there  is  a  middle  ground 
upon  which  we  should  all  unite,  namely :  First,  careful  determination  and 
description  of  worms  upon  internal  topographical  anatomy;  secondly, 
a  thorough  study  of  the  history  (both  life  history  and  bibliographic 
history)  of  each  species;  thirdly,  histological  details,  and  fourthly,  a 
consistent  application  of  such  rules  of  nomenclature  as  will  render  all 
of  our  work  international:  International  Rules. 

Rudolphi1  in  1804  originally  described  Tcenia  denticulata  as  follows : 

Unter  den  Eingeweidewiirmer  [i.  e.,  in  the  Hanover  collection]  war  ein  Stuck,  das 
mieh  sehr  interessirte,  namlich  ein  Bandwurm,  der  in  Havemanns  Gegenwart  einem 
Kalbe  abgegangen  war;  er  war  aber  ohne  Kopf.* 

*Ich  habe  diese  Wiirmer  im  Museum  zu  Alfort  wieder  gefunden,  und  auch  einige 
Exemplare  geschenkt  bekornmen.  In  seiiiem  itzt  ganz  vergrifi'enen  Werke  iiber  die 
Wiirmer  hat  Chabert  ihn  mit  alien  iibrigen  Bandwurmem,  unter  dem  Namen  Tenia 
rubna6  [ruband]  zusammengeworfen ;  im  Museum  hingegeii  war  der  Bandwurm  aus 
Darm  und  Magen  der  Kuh  ohne  Noth  in  zwey  Arten  getheilt.  Es  ist  eine  wirklich 
neue,  noch  nirgends  beschriebene  Art,  die  ich  Tcenia  denticulata  nenne,  und  an  einem 
andern  Ort  ausfiihrlich  beschreiben  werde.  Der  Wurm  ist  5  bis  16  Linien  lang,  die 
vordern  Glieder  sind  2  bis  5  Linien,  die  hintern  beynahe  einen  Zoll  breit;  der  kleine 
viereckige  Kopf  unbewaffnet,  mit  vier  in  Kreuz  stehenden  Saugmiindungen ;  kein 
Hals,  alle  Glieder  kurz,  mit  Oeffnungen  an  beyden  Seiten,  aus  denen  mehrentheils 
ein  kleiuer  spitzer  Korper  hervorsteht,  wodurch  der  Wurm  wie  gezahnt  erscheint. 
Ich  selbst  habe  ihn  nie  beym  Kindvieh  gefunden. 

Later,2  in  1805,  he  adds : 

14.  Tcenia  denticulata  mihi.    T.  ru bane's  et  lance"ole"s  dans  les  intestins  d'une  vache. 

^emerkungen  aus  dem  Gebiete  der  Naturgeschichte,  Medizin  und  Thierarzney- 
kuude  auf  einer  Reise  durch  einen  Theil  von  Deutschjand,  Holland  und  Frankreich, 
I,  Berlin,  p.  81. 

2 Bemerkungen,  etc..  II,  p.  39. 


176  TAPEWORMS  OF  HARES  A  XL)  RABBITS— STILES. 


I ni  Katalog  steht :  Deux  especes  de  Tdnias  rendues  par  la  vaclie.  Auf  eiueiu  anderu 
Glase  steht:  Tenias  rubanes  dans  la  caillette  d'une  vaclie.  Ich  muss  aber  alles  fiir 
einerley  balten,  und  liabe  scbon  vou  dem  Wurni  eine  kurze  Besckreibuug  gegeben 
wie  icb  von  dem  Kabinet  der  Thierarzneyschule  in  Hannover  sprach. 

Still  later  Eudolphi1  in  1810  continues  the  parasite  under  the  same 
name  and  discusses  it  more  in  detail.  His  entire  remarks  are  quoted 
here,  as  they  are  important,  being  based  upon  the  original  material.2 

[P.  79.]  TAENIA  J>ENTICULATA,  K.     Taenia:  capite  tetragono,  collo  nuUo,  articulis  bre- 

vissimis  foraminibus  marginalibus  oppositis,  lemuiscis  dentiformibus. 
[P.  80.]  Camper  >o  Beschiift.  Berl.  N.  Fr.,  4,  p.  139. 

Gnielin,  Syst.  Nat.,  p.  3074,  n.  55,  Taenia  ovina  ft.  bovis. 

Carlisle  Transact.  Soc.  Linn.  II,  tab.  25,  fig.  15,  16.     Taenia  ovina,  bovis. 

Rudolphi  bodoepor,  Vol.  I,  p.  81 ;  Vol.  II,  p.  39.     Taenia  denticulata. 

Hab.  in  Jlwe.  Camperus  a  bove;  Havemannus,  Scliolae  Vet.  Hannov.  Director 
meritissimiis-  a  vitulo  (solitariani,  capite  destitutam);  Cbabertus  a  vacca  copiose, 
dejectam  observaveruiit,  hie  etiani  in  vaccae  ventriculo  quarto  reperit.  Specimiua 
mea  ex  Mnseo  Scbolae  Veterinariae  Altorfensis  ditissimo. 

Descr.  Vermes  quindecim  ad  sedecim  polliceslongi,  antice  duas  ad  quinque  liueas, 
postice  frve  pollicem  lati;  coloris  albidi,  vel  grisei. 

Caput  ^xiguuin,  tetragonum,  latiusculum,  osculis  quatuor,  anticis,  subcontiguis, 
subglobosis,  liorum  apertuia  exigua,  orbiculari.  Collum  iiullum.  Articuli  aliquot 
capiti  proximl  angusti,  mox  vero  latiores,  tandem  latissimi  et  subaequales  fiunt,  ita 
tamen,  ut  in  uno  alterove  specimine  media  verinis  pars  paullo  angustiores  et  simul 
longiore^  objiciat;  omnes  ce.teroquiu  brevissimi,  ut  longitudo  latissimorum  vix 
liueam,  Tplurimorum  no  dimidiam  quidein  excedat.  Margines  articuloruni  postici 
crenati  sive  undulati  superficieni  insequentium  qua  partem  tegnnt;  latcrales  (aute- 
rioribus  nonnullis,  rarius  mediis  quibusdam  articulis  exceptis)  obtusiusculi  [p.  81], 
foramine  utrinque  medio,  opposite,  insignes,  e  quo  denticulus  acutus,  leviter  reflex  us 
et  duriu^culus  (lemniscus)  exseritur.  Substantia  mollis,  plus  minus  crassiuscula,  ut 
crassities  iuterdum  lineam  adaequet.  Ova  in  substantia  media  cumulata,  ovariis 
regularibus  mibi  lion  visis. 

Obs.  1.  Carlisle  in  figuris  citatis  canales  articulorum  laterales  longitudinem  versus 
decurrentes  et  ovaria  in  racemum  brevisainium  et  latissimum  digesta,  babet ;  baec 
omnia  lie  in  tenuioribus  quidein  speciminibus  vidi,  vel  sectione  detegere  potui. 
Denticuli  validi  liorunKiue  foramina  ab  eodem  minus  bene  sistuntur. 

Obs.  2.  Chabertus  in  opere  Bibl.  n.  19.9.3  indicate  Taeniam  hancce  cum  reliquorum 
mammalmm  Taeniis,  sub  nomine  T£nia  ruban6,  perperam  conjunxit,  in  Musei  Alfor- 
tensis  autem  catalogo  duas  in  species  (T<5nia  rubane"  et  Tenia  lanceole)  praeter  ullam 
necessitatem  divisit,  nani  specimina  niiuora  quam  reliqua  ad  formaui  lanceolatana 
magis  accedere  nil  refert. 

Obs.  3.  A  specie  praecedente  notis  in  obs.  2.  ad  eandem  indicatis  abunde  differt 
quibus  autem  Taenia  caprae  n.  84.  dicta  distinguenda  sit,  hujus  capite  nou  viso 
determinare  non  ausini;  pars  antica  tanaeji  quam  in  denticulata  tenuior,  licet  quaiu 
in  praec«dente  latior,  videtur. 

That  the  citations  from  Camper,  Carlisle,  and  Gmelin  must  be 
rejected  from  the  bibliography  of  this  species  follows  from  the  data 
given  on  page  177. 

^ntozoorum  sive  vermium  intestinalium.     Historia  naturalis,  II,  pt.  2,  p.  39. 

2The  original  description  of  1804  is  evidently  based  upon  the  Alfort  material  and 
not  upon  tlie  specimen  seen  at  Hanover.  Chabert's  specimens  must  therefore  be 
taken  as  types. 

3  Traite"  des  maladies  vermineuses  dans  les  animaux.  Paris,  1782;  second  edition, 
1787  J  German  translation,  Gottingen,  1789. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  177 

Gurlt1  in  1831  obtained  Budolphi's  specimens  and  gave  two  figures 
of  the  worm.  Creplin2  in  1842  also  studied  the  originals  and  gave 
quite  a,  fair  description  of  them  (for  details  see  Stiles3). 

Since  Creplin's  time  numerous  authors  have  mentioned  T.  denticulata 
as  a  parasite  of  cattle  and  sheep,  and  quite  a  number  of  specimens 
have  been  determined  as  belonging  to  this  species  (for  details  see 
Stiles'1).  ^  In  1891  Blanchard5  placed  this  species  in  his  genus  Moniezia. 
Later,0  in  1S!)3, 1  studied  several  end  segments  of  Kudolphi's  originals, 
compared  them  with  several  specimens  determined  as  T.  denticulata 
by  various  helminthologists,  and  called  attention  to  the  disagreement 
in  the  statements  of  different  authors  as  well  as  TO  the  fact  that  none 
of  their  specimens  agreed  with  J  udolphi's  original  type.  The  few 
statements  I  made  upon  the  originals  were  extremely  meagre  and 
guarded,  showing  that  "we  know  practically  nothing  of  the  micro- 
scopic anatomy  of  M.  denticulata." 

Upon  examining  originals  of  Baird's  Tcenia  f/oesei  and  Riehm's 
Dipylidium  latissimum,  Hassall  and  Stiles  were  surprised  to  find  the 
great  resemblance  the  cirrus  bore  to  the  cirrus  in  the  few  segments  of 
Bndolphi's  material  deposited  in  the  United  States  National  Museum. 
Through  the  kindness  of  Geheimrath  Mobius  and  Dr.  A.  Collin  we 
obtained  further  material  of  liudolphi's  originals  and  were  able  to 
prove7  that  Tcmia  denticulata  contained  two  distinct  species  of  cestode, 
i.  e.,  one  agreeing  Avith  Biehm's  I),  lattssimum  and  T.  goezcij  the  other 
with  Cteuotcmia  pectinata  (Goeze,  1782  partiin,  Biehm,  1881),  Railliet, 
1893.  We  then  expressed  the  opinion  that  an  error  had  occurred  in 
the  original  label  of  Budolphi's  specimens,  and  that  they  were  in  real- 
ity leporine  rather  than  bovine  cestodes. 

Baird's  description8  of  T.  goezei,  in  1853,  reads  as  follows: 

19.  Tcenia  Goezii,  Baird.  Head  wanting.  Articulations  of  body  very  short,  numer- 
ous. The  inferior  margin  (straight.  Genital  orifices  opposite,  situated  on  or  near 
the  lower  edge  of  each  joint,  the  lemniscus  being  projected  out  in  form  of  an  ele- 
vated papilla,  which  curves  downward.  Greatest  "breadth  of  body  6  lines,  length 
of  articulations  about  |  a  line. 

This  species  differs  from  expansa  and  denticulata  in  having  the  posterior  border  or 
edge  of  each  joint  smooth  and  rounded,  instead  of  being  crenulate  or  undulated, 
and  having  the  genital  orifices  situated  on  the  lower  edge  of  the  joint  instead  of  in 
the  middle. 

Hab.  (!) 

From  old  collection. 

1Lehrbuch  der  pathologischen  Anatomie  der  Haussiiugethiere,  p.  381,  pi.  x,  figs. 
3-4. 

2  Endozoologische  Beitrage,  Weigmanu's  Arch.  Naturg.,  I,  pp.  315-327. 

3  Revision  of  adult  Cestodes  of  cattle,  sheep,  and  allied  animals,  Bull.  4,  Bur.  An. 
Ind.,  U.  S.  Dept.  Agric.,  1893,  p.  44. 

4  Loc.  cit.,  1893,  pp.  44-46. 

6Me"m.  Soc.  zool.  France,  IV,  1891,  p.  187. 

6  Loc.  cit.,  1893,  pp.  42-47. 

7  Vet.  Mag.,  Ill,  1896,  p.  6-9. 

8  Cat.  Specs.  Ent.  or  Intestinal  Worms,  Coll.  Brit.  Mus.,  Lond.,  p.  78. 

Proc.  K  M.  vol.  xix 12 


178  TAPEWORMS  OF  HA E ES  A XD  E A B BTTS—S TILES.  VOL. xix. 


Riehm1  in  1881  studied  the  tapeworms  of  rabbits  and  hares,  and  in 
a  preliminary  account  of  his  work  described  this  same  form  as  Citto- 
tcenia  latissima,  new  genus,  new  species.  Later2  he  placed  this  species 
in  the  genus  Dipylidium  with  the  following  specific  diagnosis : 

Kopf  hakenlos  liber  f  min.  breit,  rnit  stark  vorspringendeu  Saugnapfen  und 
da<lurch  gegen  die  lanzettformig  sich  verbreitermlo  Gliederkette  deutlich  abgesetzt. 
Geschlechtsciffnungen  beiderseits,  in  den  zitzenartig  vorspringenden  Hinterecken 
£er  Glieder,  welche  den  Riindern,  besonders  der  contraliierten  Thicre  ein  gefranztes 
Ansehen  verleihen.  Glieder  stets  viel  kiirzer  als  breit  und  uamontlich  nach  den 
Seiten  bin  ungemein  dick.  Liinge  im  gestreckten  Zustaude  bis  80  cm.,  Breite  der 
reifsten  Glieder  15  mm.  und  dariiber  Wobnthier:  Lepus  cuniculus. 

Of  anatomical  details  he  gives  the  following,  which  appear  to  me  to 
be  of  importance: 

Segments  may  attain  3  to  3.5  mm.  in  thickness;  the  strobila  is  generally  found  in 
the  lower  portion  of  tbe  small  intestines  and  is  of  a  grayisb  to  a  reddish  gray 
in  color,  something  like  Lothrioceplialtis  latns.  Black  pigment  on  the  suckers  and 
genital  pores.  Suckers  prominent;  neck  short.  Genital  anlagen  appear  very  early. 
Male  organs:  Testicles  very  numerous,  0.115  mm.  in  diameter,  scattered  throughout 
dorsal  portion  of  median  field;  cirrus  pouch  large,  provided  with  three  layers  of 
mucles,  two  circular  and  one  longitudinal  layer,  cirrus  is  generally  protruded. 
Female  organs :  Vagina  opens  close  to  cirrus,  and  is  quite  thin;  median  to  the  longi- 
tudinal canals  it  swells  into  a  receptaculum  seuiinis;  female  glands  essentially  the 
same  as  in  Dipylidium  pectinalitm,  but  situated  very  close  to  the  longitudinal  canals. 
The  uterus  is  described  "'not  as  a  simple  tube,  widened  here  and  there,  and  running 
transversely  through  the  segment,  but  it  is  broken  up  into  two  or  three  such  tubes, 
which  unite  here  and  there,  and  thus  cut  the  parenchyma  up  into  islands.  This 
structure  is  seen,  however,  only  in  segments  without  eggs.  Through  the  pressure  of 
the  developing  ova  these  tubes  widm  so  that  the  islands  of  parenchyma  are  sup- 
pressed and  the  uterus  then  appears  as  a  single  tube."  In  some  segments  an  extra 
(third)  set  of  female  glands  was  observed  right  or  left  of  the  median  line.  Excre- 
tory system  very  highly  developed;  in  the  younger  segments  it  is  composed  of  a  net- 
work of  canals,  with  one  lateral  canal  which  is  especially  large;  transverse  canal 
with  numerous  anastomoses  present;  as  the  genital  organs  develop,  the  canals  of  the 
median  field  become  more  or  less  suppressed,  but  three  or  four  lateral  canals  persist 
each  side  and  the  transverse  canals  at  the  posterior  margin  become  larger,  so  that 
the  excretory  system  now  bears  more  of  a  resemblance  to  that  of  JJipylidium  pectina- 
tum;  the  second  canal  system  (dorsal  canal)  resembles  that  of  Dipylidium  leiickarti. 
(Abstract.) 

Neumann3  in  1888,  places  this  worm  in  the  genus  Tcenia. 

Blanchard4m  1891  examined  Baird's  specimens  and  recognized  them 
as  identical  with  D.  latissimiim,  Eiehm.  He  also  examined  fresh  speci- 
mens, which  he  describes  in  some  detail. 

Strobila  measures  40  to  50  cm.  long  and  contains  about  210  segments ;  bead,  0.8  mm. 
broad  by  0.63  long.  Neck  absent,  segments  may  attain  4  mm.  in  length  by  15  in 
breadth.  Genital  pore  appears  about  the  seventy-eighth  segment;  in  the  older  seg- 
mnets  it  is  more  distal  than  in  the  younger  segments ;  egg  is  spherical  52  to  60  //; 
bulb  of  pyriform  body  16  to  24  //,  horns  long  and  curved.  (Abstract.) 

1  [Untersuchungen  an  den  Bandwiirmern  der  Hasen  und  Kaninchen,  Zeitschr.  ges. 
Naturw.  3  ser.,  VI,  1881,  p.  200. 

2Studieu  an  Cestodeu,  Zeitschr.  ges.  Xaturw.,  3  ser.,  VI,  1881,  pp.  583-590. 

3Trait6  des  maladies  parasitaires,  p.  426,  Paris,  1888. 

4Me"m.  Soc.  zool.  France,  IV,  1891,  pp.  444,  451,  452-457,  figs.  21-25. 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUV.  179 

Bailliet1  iu  1893  places  Riehm's  species  in  his  genus  Ctetiotcmiin7~ 

From  the  work  of  former  authors  and  from  my  own  observations  upon 
the  original  material  of  T.  dentieulata,  T.  (joezei,  and  Clttotcenia  latis- 
sima,  I  propose  the  following  as  a  revised  specific  diagnosis: 

Diagnosis.— Cittotcmia  dentieulata  (Kudolphi,  1804),  Stiles  and  Has- 
sall,  189(5:  Strobila  attains  40  to  50  cm.,  even  80  cm.  in  length  by  15  mm. 
in  breadth  by  3.5  mm.  in  thickness,  is  grayish  to  grayish  red  in  color, 
and  is  made  up  of  200  or  more  segments.  Head  unarmed,  0.8  mm! 
broad  by  0.63  mm.  long,  suckers  large  and  prominent;  rostellum  not 
observed,  Neck  ub> ent.  Genital  pores  double,  prominent,  opposite  in 
posterior  half  of  margin,  near  corner  of  the  segment.  Male  organs: 
Testicles  0.115  mm.  in  diameter,  very  numerous,  scattered  through  the 
dorsal  portion  of  the  median  field;  cirrus-pouch  large  and  prominent, 
1.12  mm.  long  by  0.32  mm.  broad.  Female  organs:  Vagina  runs  ven- 
trally  and  distally  of  cirrus-pouch  on  both  sides  of  segment;  female 
glands  very  similar  to  those  of  Gittotcenia  pcctinata,  situated  close  to 
longitudinal  canals ;  uterus  very  complex.  Eggs  52  to  63  //in  diameter; 
bulb  of  pyriform  body  16  to  24  /--.  Excretory  system :  Very  complex, 
numerous,  thin-walled  longitudinal  canals  (probably  all  belonging  to 
the  ventral  system);  dorsal  canals  with  thick  wall.  Genital  canals 
cross  the  longitudinal  canals  and  nerves  dorsally. 

Host. — Common  European  wild  rabbit  (Lepus  cuniculus). 

Types.— Eudolphi's  type  in  Berlin  Museum;  fragments  in  the  collec- 
tion of  Stiles,  Nos.  17, 1492,  U.S.  <ST.M.  Baird's  type  in  South  Kensing- 
ton Museum  (London);  fragments  in  collection  of  Blanchard.  Riehm's 
cotypes  in  collection  of  Leuckart;  Vienna  Museum;  Xo.  1328,  U.S.N.M. 
Several  typical  specimens  in  collection  of  Moniez. 

Geographical  distribution. — Europe:  France  (Chabert,  E.  Blanchard, 
Monie/);  Saxony  (Biehm). 

CITTOT^NIA  CTENOIDES  (Railliet,  1890),  Stiles,  1896. 
(Plate  XIV,  figs.  1-8.) 

1881,  DipyMium  leuckarti,  RIKHM,  Zeitschr.  f.  d.  ges.  Xaturw.,  3  ser.,  VI,  p.  200;  pp. 

566-577,  pis.  v,  figs.  3, 11-13, 16;  vi,  figs.  5-6. 
1888,  Tasnia  leucJcarti  (RiEiiM,  1881),  NEUMANN,  Traite  des  maladies  parasitaires, 

etc.,  p.  426  [ncc  Tccnia  leuckarti,  KRABBE,  1869,  Bidrag,  p.  335]. 

1890,  T.  ctenoides,  RAILLIET,  Rev.  d.  Sci.  nat.  appliq.  Bull.  Soc.  nat.  d'Acclimat.  de 

France,  p.  346. 

1891,  Moniezia  huckarti    (RIEHM,  1881),  R.  I^LAXCHARD,  Mdm.    Soc.   zool.,  France, 

IV,  p.  187;  pp.  444,  450-451. 
1893,  Ctenotwnia  Jeuckarti  (RiEHM,  1881),  RAILLIET,  Traitd  de  Zool.  med.  et  agric., 

I,  p.  278. 
1896,   Cittotocnia  leuckarU  (RIEHM,  1881),  STILES  &  HASSALL,  Veterinary  Magazine, 

III,  p.  407. 

For  the  history  of  this  species  prior  to  Riehm,  1881,  see  the  historical 
review  of  G.  pectinata,  p.  184. 


Traito  Zool.  med.  et  agric.,  I,  p.  278. 


ISO  TAPEWORMS  OF  HARES  AND  RABBITS—  STILES.  VOL.XIX. 

Eiehm1  in  188  L  was  the  tirst  to  clearly  define  this  form,  which  he 
named  Dipylid'nun-  Icuckarti,  with  the  following'  specific  diagnosis: 

Kopf  hakeulos,  sehr  klein,  etwa  0.5  mm.  breit,  mit  4  nachaufliegenden  Sang- 
napfen,  nicht  abgesetzt  gegen  den  Hals,  welcher  mit  lanzettformiger  Verbreiterung 
in  die  Gliederkette  iibergeht.  Geschlechtsoffnungeii  beiderseits  im  hinteren  Vier- 
theil  des  Proglottisrandes.  Glieder  trapezforrnig,  aueh  im  gestrecktesten  Zustande 
noch  breit,  fiir  gewohnlich  aber  etwa  3-6  rnal  breiter  als  lung.  L-inge  im  ges- 
treckten  Zustande  bis  80  cm.,  Breite  der  reifsten  Glieder  bis  1  em.  Die  Strobila 
hliufig  durch  Liingsfalten  der  Haut  wie  gestreift.  AVohiithier  :  Lepus  cinriculu*. 

Iii  the  anatomical  discussion,  and  in  his  figures,  Eiehm  brings  out 
the  following  points  : 

The  head  measures  0.5  mm.  broad  by  about  0.166  mm.  thick;  neck  moderately  long, 
its  distal  end  nearly  twice  as  broad  as  its  proximal  end  ;  500  to  750  proglottids  present 
in  the  strobila  ;  posterior  edge  of  segments  extends  prominently  over  the  next  follow- 
ing segment,  so  that  the  side  of  strobila  appears  serrate.  Male  organs  :  Testicles  50 
to  60  f.i  in  diameter,  arranged  in  two  groups  in  each  segment,  one  on  each  side  of  the 
median  field  near  the  ovary  ;  cirrus  pouch  small,  sacular,  contains  a  small  vesicula 
semiualis  and  the  retracted  and  coiled  penis.  Female  organs  :  The  female  anlageu 
appear  as  early  as  the  twenty-fifth  segment;  female  glands  quite  close  to  ventral 
canal  ;  uterus  same  as  in  C.  pectinata.  Excretory  system  :  Circular  canal  said  to  be 
absent  from  the  head;  a  median  canal  forms  a  dorsal  and  a  ventral  loop  between  the 
suckers;  each  loop  branches,  one  forming  the  two  ventral  canals,  the  other  the  two 
dorsal  canals;  the  ventral  canals  are  connected  by  the  transverse  canals  in  each 
segment,  but  in  the  posterior  segments  a  network  of  canals  supplants  the  ordinary 
transverse  canals;  a  second  canal  system  —  evidently  tho  dorsal  canals  —  is  described 
as  possessing  a  thick  muscular  layer.  It  is  figured  as  lying  between  the  ventral 
canal  and  nerve.  Ova,  67  to  70  JLI  in  diameter.  Of  more  than  150  wild  rabbits,  only 
10  to  12  were  not  infested  with  this  parasite.  (Abstract.; 

Neumann2  in  1888  placed  the  worm  in  the  genus  Tcenia;  later"  he 
reverted  to  Dipylidium  and  recorded  a  worm  from  the  domesticated  rab 
bit  which  he  considered  identical  with  this  form.     Eailliet  in   1890 
found  this  worm  in  France  and  proposed  the  name  Tcenia  ctenoides  in 
place  of  T.  leuckarli  (Eiehm,  1881),  Neumann,  1888  [nee  Krabbe,  I860]. 

K.  Blanchard4  in  1891  placed  this  species  in  the  genus  ILoniezia, 
translating  Eiehm's  diagnosis. 

Eailliet  in  1893  transferred  the  species  to  his  genus  Ctenotcenia. 

I  have  examined  one  of  Eiehm's  originals,  and,  through  the  kindness 
of  E.  Blauchard,  I  have  received  several  tapeworms  under  the  label 
u  Tcenia  pectinata^  Lapin  domestique,  10,  8,  89.  G.  Marclial.  Hucque- 
liers  pres  Montrei  —  [?]  sur  mer."  These  worms  agree  with  Eiehm's 
specimens  of  Dipylidium  leuckarti.  One  of  Moniez's  specimens  labeled 
"Azores"  is  also  a  D.  leuckarti.  Upon  the  material  at  my  disposal  and 
Eiehm's  anatomical  discussion  I  base  the  following  revised  specific 
diagnosis  : 


.  an  Cestoden.     Zeitschr.  ges.  Naturw.,  3  ser.,  VI,  p.  200;  pp.  566-575. 
2  Traite  des  maladies  parasitaires,  Paris,  1888,  p.  426. 

3Trait6  des  maladies  parasitaires,  Paris,  2d  ed.,  1892,  p.  461.    Parasites  and  Parasitic 
Diseases  of  Domesticated  Animals,  London,  1892,  p.  473. 
4  Mem.  Soc.  zool.  France,  IV,  p.  187;  pp.  450-451. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEl'M. 


Diagnosis.— Cittotcenia  ctenoid™  (Kailliet,  1800),  Stiles,  1896.  Stro- 
bila  serrate  may  attain  80  cm.  in  length  by  10  mm.  in  breadth ;  composed 
of  500  to  750  segments,  which  are  always  at  least  3  to  5  times  broader 
than  long.  Head  unarmed,  about  0.44  to  0.5  mm.  broad  by  0.16  mm. 
thick;  no  rostelluin  visible.  Suckers  0.176  mm.  in  diameter.  Neuk 
short  and  broad,  or  absent,  segmentation  beginning  immediately  back 
of  the  head.  Genital  pores  double,  situated  in  posterior  half  of  margin. 
Male  organs:  Testicles  arranged  in  two  groups  in  lateral  portion  of 
median  field;  about  60  to  80  testicles  to  each  group,  50  to  60  it  in 
diameter.  Cirrus  pouch  small,  0.16  mm.  long  by  0.128  mm.  broad;  it 
does  not  reach  the  longitudinal  nerve;  contains  the  coiled  and  inverted 
cirrus  and  a  small  vesicula  seminalis.  Female  organs:  Ovary,  vitello- 
gene,  and  shell  glands  agree  in  general  with  the  same  organs  in  C. 

ctinatajbutare  situated  much  closer  to  the  longitudinal  canals;  uterus 
agrees  with  that  of  C.pectinata,  but  appears  to  be  rather  more  simple. 
Dorsal  canal  lies  between  ventral  canal  and  nerve  or  dorsal  of  ventral 
canal.  Vas  deferens,  vagina,  and  uterus  extend  from  median  field  into 
lateral  field  dorsally  of  the  longitudinal  canals  and  nerves.  Ova,  64  ^ 
in  diameter;  bulb  of  pyriform  body,  20  /,/. 

Hosts.—  European  wild  rabbit  (Lepus  cuniculus],  by  Riehm;  tame 
rabbit  (L.  cuniculus  domesticm),  by  Neumann,  Railliet,  and  Marchal. 

Types.— Cotype,  No.  1327,  U.S.N.M.  (from  collection  of  Riehm). 
Typical  specimens:  No.  114,  U.S.N.M.  (from  collection  of  Blanchard); 
Nos.  1449.4  and  1457,  U.S.N.M.  (from  collection  of  Moniez);  collection 
of  Blanchard;  collection  of  Moniez. 

Geographical  distribution. — Europe:  Saxony,  by  Riehm;  France,  by 
Neumann,  Railliet,  Marchal,  Moniez.  The  Azores  (collection  of  Moniez). 

CITTOT^NIA   PR^COQUIS  (Stiles,  1895),  Stiles  &  Hassall,  1896. 
(Plate  XV,  figs.  1-6.) 

1895,  Ctenotcenia  prcecoquus  [typographical  error  for  prcecoquis],  STILES,  Veterinary 

Magazine,  II,  June,  p.  345.     Aug.  28, 1895. 

1896,  Cittotcenia  prwcoquis  (STILES,  1895),  STILES  &  HASSALL,  Veterinary  Magazine, 

III,  p.  407. 

Prof.  Herbert  Osborne,  of  Ames,  Iowa,  collected  two  specimens  of 
tapeworms  from  the  Pocket-Gopher  (Geomys  bursarius],  which  he  pre- 
sented to  the  Bureau  of  Animal  Industry.  The  species  is  rather  a 
difficult  one  to  interpret,  especially  upon  press  preparations,  and  the 
limited  amount  of  material  does  not  allow  a  minute  anatomical  analysis 
by  means  of  sections,  but  sufficient  characters  can  be  given  to  enable 
one  to  easily  recognize  the  form. 

The  specimens  measure  about  40  mm.  long  and  contain  about  150  seg- 
ments ;  the  greatest  breadth,  about  5.5  mm.,  is  reached  about  10  mm.  from, 
the  distal  extremity;  the  distal  10  mm.  becomes  narrower.  Head  not  very 
ilistinct  from  the  strobila.  0.43  nun.  broad  by  0.32  mm.  long.  No  rostel- 
lum  visible;  no  hooks  present;  sticker  0.16  mm.  long  by  0.128  mm.  broad. 


182  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 


Neck  very  short  and  broad.  Segmentation  visible  0.5G  mm.  from  ante- 
rior extremity  of  the  head.  The  proglottids  rapidly  become  distinct,  and 
in  the  specimens  at  hand  the  posterior  border  of  each  segment  overlaps 
the  anterior  border  of  the  next  following  segment  for  about  one-third 
of  its  length  in  the  anterior  segments  to  one-sixth  of  its  length  in  the 
posterior  segments.  The  genital  aiilagen  become  visible  almost  imme- 
diately after  the  segmentation.  The  genital  pores  are  double  and 
opposite  in  about  the  middle  of  the  lateral  margin.  Male  organs :  The 
testicles  are  numerous  and  are  scattered  over  the  entire  dorsal  portion 
of  the  median  field,  but  do  not  quite  reach  the  lateral  canals;  in  tbe 
middle  portion  of  the  median  field  they  extend  farther  ventral  than  in 
the  lateral  portion.  The  cirrus  pouch  lies  dorsal  and  slightly  posterior 
to  the  vagina,  on  both  sides  of  the  segment;  it  is  0.24  mm.  long  by  96^ 
broad  (proximal  end) ;  it  is  relatively  muscular;  the  lateral  portion  is 
narrow,  but  the  proximal  portion  is  swollen  to  contain  a  globular 
vesicula  seminalis;  the  vas  deferens  coils  and  curves  from  here  median 
and  cephalad,  the  convexity  of  the  curve  being  distal.  Female  organs : 
The  female  glands  lie  in  the  median  field,  about  halfway  between  the 
median  line  and  lateral  canals ;  they  were  not  analyzed  in  detail ;  details 
of  the  development  of  the  uterus  must  also  be  left  to  some  one  who  can 
obtain  further  material;  the  vagina  lies  ventral  and  slightly  proximal 
to  the  cirrus-pouch  on  both  sides  of  the  segment;  it  is  narrow,  19  mm. 
long,  and  extends  to  a  point  slightly  dorso-median  of  the  ventral  canal, 
where  it  swells  into  an  enormous  receptaculum  seminis  which  occupies 
the  ventro-lateral  portion  of  the  median  field,  and  in  some  sections 
measures  0.72  mm.  long  by  0.19  mm.  broad;  median  of  this  organ  and 
ventral  of  the  testicles  are  situated  the  female  glands;  there  is  appar- 
ently a  common  uterus  to  both  ovaries;  it  is  situated  vent-rally  of  the 
testicles  at  the  height  of  the  genital  pores,  but  proximally  and  distal  ly 
of  this  plane  it  extends  farther  dorsad:  sections  on  other  planes  than 
that  of  the  pore  show  that  dorso- ventral  trabecuhe  of  the  body  stroma 
divide  the  uterus  into  numerous  pouches,  although  sections  at  the 
pores  generally  show  a  more  or  less  continuous  transverse  stem;  the 
ova  are  unusually  small,  measuring  32  to  36  //  in  diameter;  the  bulb 
of  the  pyrifonn  body  varies  between  8  //  and  12  //;  the  horns  are 
relatively  long  and  crossed.  Excretory  and  nervous  systems:  At  the 
height  of  the  genital  pores  the  dorsal  canal  lies  between  the  ventral 
canal  and  nerve;  in  other  planes  it  may  lie  dorsal  of  the  ventral  canal. 
The  genital  canals  pass  dorsally  while  the  uterus  passes  vent-rally  of 
the  longitudinal  canals  and  nerves. 

As  specific  name,  I  proposed  prwcoquis  (misprinted  as  prwcoquus  in 
the  original  description),  referring  to  the  unusually  early  development 
of  the  genital  organs.  The  following  is  proposed  as  a  specific  diagnosis : 

Diagnosis. — Cittotcenia  prcccoqiiis  (Stiles,  1895),  Stiles  &  Hassall, 
189G.  Strobila  attains  about  40  mm.  in  length  by  5.5  mm.  in  breadth 
by  0.72  mm.  in  thickness;  the  posterior  10  mm.  is  slightly  narrowed. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  183 

Head  unarmed,  not  very  distinct  from  strobila,  0.43  mm.  broad  by 
0.32  mm.  long ;  rostellum  not  observed  5  suckers  0.16  mm.  long  by 
0.128  mm.  broad.  Neck  short  and  broad.  Segmentation  begins  about 
0.56  mm.  from  the  anterior  extremity  of  head.  About  150  proglottids 
present,  very  distinct;  the  posterior  flap  overlaps  the  anterior  one-third 
to  one  sixth  of  the  next  following  segment.  Genital  pores  double  and 
opposite,  not  prominent,  in  about  the  middle  of  the  lateral  margin. 
Male  organs:  Testicles  numerous,  confined  to  dorsal  portion  of  median 
field;  cirrus  pouch  dorsal  of  vagina  ou  both  sides  of  segment,  0.24  mm. 
long,  muscular,  containing  globular  receptaculum  seminis  (about  90  ^ 
diameter)  in  its  proximal  portion;  it  extends  to  or  slightly  beyond  the 
ventral  canal.  Female  organs:  Female  glands  about  halfway  between 
median  line  and  ventral  canal;  vagina  narrow,  extends  across  ventral 
canal,  then  dilates  into  a  large  receptaculum  seminis;  uterus  probably 
single,  possessing  blind  proximal  and  distal  pouches.  Ova,  32  to  36  u 
in  diameter;  bulb  of  pyriform  body,  8  to  12  yu;  horns  long  and  crossed. 
Excretory  system :  Dorsal  canal  between  ventral  canal  and  nerve  at 
height  of  the  genital  canals;  genital  canals  cross  dorsally,  but  uterus 
crosses  ventrally,  of  the  longitudinal  canals  and  nerves. 

Hosts. — Pocket  Gopher  (}Gcomys  bursarius  (Shaw,  1800)  ),  collected 
by  Herbert  Osborne  at  Ames,  Iowa. 

Types. — Bureau  of  Animal  Industry  cestode  series  No.  1079  desig- 
nated as  type  and  deposited  in  the  United  States  National  Museum; 
paratype  (1372,  part  in  sections)  deposited  with  type.  Technique  of 
types :  Corrosive  sublimate,  alcohol,  acid  carmine. 

PECTINATA  GROUP. 

The  European  form  Cittotcenia  pectinata  forms,  with  certain  American 
parasites,  a  very  compact  group,  but  it  is  difficult  to  know  what  rank 
should  be  given  to  this  division.  Generic  rank  would  be  too  high, 
specific  rank  too  low.  I  propose  to  include  these  cestodes  in  the  Pec- 
tinata  group,  which  I  base  upon  the  peculiar  cirrus  pouch. 

Three  species  come  within  this  group  at  present: 

1.  Type  species  C.  pectinata,  as  defined  below,  p.  188,  its  chief  char- 
acters being  the  length  of  the  cirrus  pouch,  the  quadrangular  arrange- 
ment of  the  testicles,  and  their  extension  beyond  the  ovaries  to  the 
lateral  canals. 


'Synonymy  after  Merriam:  1800,  M«s  bursarins,  Shaw;  1815,?  M.  ludoricianus, 
Ordway ; *1817,  Diplostoma  fusca,  Rafinesque;  1817,  Geomtj8  cinerea,  Rafinesque  (Mas 
ftMrartrmsivnamed);  1820,  Saccophoru*  bursarius,  Kuhl ;  1*21,  Mas  saccalm,  Mitcbill; 
1823,  Psetidostoma  burmrius,  Say;  (1822)  1825,  Ascomt/s  canadeiisis,  Lichteufel  1;  1S59> 
Geomi/sf  bnrsariua,  Richardson;  1852,  Geomys  canadensis,  LeConte;  1852,  G.orcyonen- 
sis,  LeConte. 

C.  Hart  Merriam,  1895.  Monographic  Revision  of  the  Pocket  Gophers,  Family 
Geomyidje  (exclusive  of  the  species  of  Thomomus),  p.  120.  North  American  Fauna, 
No.  8,  Division  of  Ornithology  and  Mammalogy,  United  States  Department  of  Agri- 
culture. 


184  TAPEWORMS  OF  HAKES  AXD  RABBITS— STILES. 


2.  The  second  species,  C.  perplexa,  is  directly  intermediate  between 
the  first  and  third,  "but  I  am  unable  to  find  connecting  links  between  it 
and  the  other  forms.     The  cirrus  pouch  is  about  two  thirds  as  large  as 
that  of  C.  pectinata  and  somewhat  more  distinct  than  that  of  C.  rarl- 
abilis;  the  testicles  are  arranged  in  two  triangles  and  extend  laterally 
beyond  the  ovaries  to  the  lateral  canals.     In  general  configuration  of 
the  body  it  resembles  C.  pectinata  very  closely,  but  is  much  shorter. 

3.  C.  variabilis  represents  the  last  form  of  the  series,  presenting  a 
cirrus  pouch  slightly  smaller  than  that  of  G.perplcxa  and  testicles  in  a 
quadrangle  which  is  confined  entirely  to  the  space  between  the  ovaries. 
It  occurs  in  three  different  forms  as  follows: 

(a)  C.  variabilis  found  in  Lepus  sylvaticus  with  characters  described 
below,  p.  1927  measuring  up  to  10  mm.  broad,  the  segments  always  much 
broader  than  long. 

(b)  C.  variabilis  angusta  differs  from  C.  variabilis  only  in  point  of  size; 
it  measures  only  2  mm.  broad  and  the  segments  are  about  three  times 
as  broad  as  long.     This  form  I  have  never  found  with  ova,  and  it  may 
be  a  case  of  arrested  development.    Those  authors  who  take  external 
form  as  a  specific  character  would  be  obliged  to  give  this  parasite  spe 
cific  rank,  but  I  am  unable  to  admit  this  view.    This  variety  is  found 
in  Lepus  sylvaticus. 

(c)  C.  variabilis  imbricata  is  the  third  variety,  and  is  one  due  proba- 
bly entirely  to  host  influence.     It  is  found  in  Lepus  palustris  in  Florida. 
The  differences  between  this  form  and  the  type  are  but  very  slight ;  the 
posterior  margins  are  more  scalloped,  the  segments  more  imbricate,  and 
the  cirrus  pouch  slightly  more  distinct. 

CITTOT^ENIA   PECTINATA  (Goeze,  1782  partim,  Riehm,  1881),  Stiles  &  Has- 

sall,  1896. 

(Plate  XVI,  figs.  1-2;  Plate  XVII,  figs.  1-2.) 

?1781,  Tcenia  acutissima,  PALLAS,  Neue  nord.  Beytrage,  I,  pp.  75-81,  pi.  in,  fig.  25. 

1<S^,  T.  pectinata,  GOEZE,  Versuch  einer  Naturgeschichte  d.  Eingeweidew.,  pp.  363- 
368,  pi.  xxvn,  figs.  7-13. 

?1800,  Alyselminthus  pectinatus  (GOEZE,  1782),  ZEDER,  Erster  Xachtrag  zur  Naturg. 
d.  Eiugeweidewurrner,  pp.  246-249. 

?1803,  Halysis  pectinata  (GoEZE,  1782),  ZEDER,  Anleitung  zur  Naturg.  d.  Einge- 
weidewiirmer,  p.  332. 

1810,  Tamia  leporiua*  RUDOLPHI  (in  synonymy),  Ent.  Hist.  Nat.,  II,  Pt.  2,  p.  82. 

1881,  Dipylidium  pectinaliim  (GOEZE,  1782  partim),  KIEHM,  Zeitschr.  f.  d.  ues. 
Naturwiss.,  3  ser.,  VI,  p.  200;  pp.  575-583,  pi.  v,  tigs.  4,  14;  vi.  iigs  4,  7. 

1891,  Moniezia  pectinata  (GOEZE,  1782  partim,  Riehm,  1881),  R.  BLAXCHARD,  ^lom. 
Soc.  zoel.  France,  IV,  pp.  187,  445,  450-452,  457-460,  figs.  26-30. 

1893,  Ctenotcenia  pectinata  (GoEZE,  1782  partim,  Riehm,  1881),  RAILLIET,  Traite  de 
Zool.  uicd.  et  agric.,  I,  pp.  278-279. 

1896,  Cittotccnia  pectinata  (GOEZE,  1782  partim,  RIEHM,  1881),  STILES  &  HASSALL,  Vet- 
erinary Magazine,  III,  p.  407. 

'Blaiichard  (1891A,  p.  457)  has  already  shown  that  Limbonrg  (1766)  did  not  use 
" Twnia  leporina"  as  a  specific  name;  the  name  therefore  dates  from  Rudolphi,  1810. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  185 

Marigues1  in  1778  mentions  and  figures  tapeworms  found  in  the 
body  cavity  of  rabbits;  but  it  is  impossible  to  determine  what  species 
he  had  before  him. 

Pallas  in  1781  found  no  tapeworms  in  the  many  hares  he  examined. 
He  received  some  worms  taken  by  Graf  von  Bork  from  hares  and 
described  them  as  Tcenia  acutissima.  The  worm  is  evidently  a  double- 
pored  leporine  form,  and  on  that  account  probably  a  CUtotccnia;  but 
from  the  description  given  by  Pallas  it  is  impossible  to  definitely  decide 
which  of  the  three  leporine  double-pored  forms  he  had  before  him. 
On  this  account  I  do  not  consider  myself  justified  in  reinstating  the 
specific  name  acutissima. 

Pallas  evidently  thought  that  Drepanidotcenia  lanceolata  of  geese 
was  identical  with  the  leporine  form,  and  he  also  placed  Schistocephalus 
from  the  body  cavity  of  Gasterosteus  side  by  side  with  the  parasite  of 
hares,  although  his  text  clearly  shows  that  he  thought  further  investi- 
gation might  result  in  separating  Schistocephalus  as  a  distinct  species. 

Goeze  in  1782  states  that  Tcenia  pectinata  is  found  in  hares  and  wild 
rabbits,  but  he  never  found  it  in  tame  rabbits;  he  mentions  its  simi- 
larity to  a  parasite  (Drepanidotcenia  lanceolata}  of  geese,  to  one  (Schis- 
tocephalus solidus)  of  the  Stickle-back  (Gasterosteus)  and  to  the  broad 
tapeworm  (Moniezia  expansa)  of  sheep,  but  does  not  consider  it  specific- 
ally identical  with  any  of  these  forms.  Goeze  found  the  species  nearly 
always  present  in  1-year  old  hares  and  in  wild  rabbits,  often  20  to  30 
in  an  individual  host.  "The  margins  of  each  segment  were  rather 
bluntly  rounded,  and  on  this  rounding,  by  means  of  the  hand  lens  a 
very  small  opening  was  visible,  out  of  which  eggs  could  be  pressed, 
and  out  of  which,  when  the  live  worms  were  placed  in  hot  water, 
through  the  sudden  contraction  of  the  canals,  they 'were  pressed  in 
small  threads."  The  first  part  of  this  quotation,  "  the  margins,  etc." 
("  Die  Seitenrande  jedes  Gliedes  kulpichtrund,  und  auf  dieser  Ruen- 
dung")  is  the  only  portion  of  Goeze's  article  which  can  be  interpreted 
as  referring  to  double  pores.  His  figures  do  not  show  pores,  but  Figure 
7  makes  the  impression  upon  a  worker  of  l)eing  a  double-pored  cestode. 
The  head  is  described  as  very  small;  segmentation  begins  almost 
immediately  back  of  the  head;  the  segments  increase  in  size  very  rap- 
idly; four  suckers  on  head,  but  no  hooks;  of  thirteen  specimens  the 
longest  measured  about  6  inches;  it  possessed  204  segments. 

Zeder  in  1800  placed  Goeze's  specific  name  in  the  genus  Alyselmin- 
thus,  and  gave  a  discussion  of  the  parasite  which  he  supposed  was 
identical  with  Goeze's  form.  He  states,  however,  that  he  found  only  a 
single  pore  to  each  segment.  While  it  seems  possible  that  Zeder's 
form  is  identical  with  Andrya  rhopalocephala,  as  assumed  by  several 
authors,  the  point  can  not  be  definitely  settled,  as  Zeder's  description 

1  Observations  sur  des  vers  Tonia  trouves  dans  le  ventre  de  quelques  lapins  sau- 
vages ;  Observations  sur  la  physique,  etc.  (Rozier),  XII,  Paris,  pp.  229-231,  pi.  n,  ng.  3. 


186  TAPE  WORMS  OF  HA  £ES  A  XD  I?  ABB  I TS— S  TILES.  VOL.  xix. 

is  too  indefinite  to  allow  a  specific  or  even  generic  determination.  In 
1803  he  transferred  the  species  to  Halt/sis. 

Eudolphi l  in  1810,  as  Eiehni  remarks,  certainly  had  more  than  Goeze's 
species  in  mind  when  he  wrote  his  diagnosis;  a  portion  of  his  descrip- 
tion applies  very  well  to  C.  denticulata,  and  a  portion  to  C.  ctenoides. 
He  includes  Zeder's  single-pored  form  in  T.  pectinata.  The  fragment  of 
Eudolphi's  (1810)  specimen  of  T.  pectinata  which  I  examined  is  so 
poorly  \ (reserved  that  no  statements  can  be  made  on  it,  but.  as  Stiles 
and  Hassall  in  1896  have  shown,  part  of  Eudolphi's  original  material 
of  T.  denticulata  belongs  to  this  species. 

Bremser 2  in  1824  gives  two  figures  of  T.  pectinata,  in  regard  to  which 
Eiehm  remarks  that  Figure  5  is  an  unquestionable  I),  leuckarti,  while 
Figure  G  (head)  is  similar  to  D.  latissimum  (=C.  denticulata}. 

Diesing's3  description  of  1850  can  be  made  to  apply  to  several  dif- 
ferent forms;  he  includes  all  the  literature  given  for  T. pectinata. 

Our  chief  knowledge  of  C.  pectinata  we  owe  to  Eiehm  in  1881,  who 
studied  its  anatomy  in  detail.  Eiehm  divided  the  heterogeneous  mass 
of  tapeworms,  which  earlier  authors  had  included  under  the  term 
T.  pectinata^  into  five  species,  which  he  named  T.  rhopalocephala  and 
T.  rhopaliocephala  (single-pored  forms),  and  Dipylidium  pectinatum,  D. 
leuckarti,  and  D.  latissimum  (double-pored  forms).  This  is  the  first 
time,  therefore,  that  T.  pectinata  was  described  in  detail,  so  that  this 
species  stands  to-day  upon  Riehm's  division,  he  having  designated  the 
particular  parasite  which  should  bear  Goeze's  specific  name. 

Eiehm 4  diagnoses  his  form  as  follows : 

Kopf  hakenlos,  ausserordentlich  kleiu,  kaum  £  inni.  breit,  gegen  die  lanzettfor- 
mig  sicli  verbreiterndo  Strobila  nicht  abgesetzt.  Geschleehtsorthungen  beidei>eits, 
fast  in  der  Mitte  des  Proglottidenrandes.  Glieder  kurz,  trapezforrnig,  anch  im 
gestrecktesten  Zustande  miudestens  4  mal  breiter  als  lang.  Liiiige  des  ansgestreck- 
ten  Wurmes  niclit  iiber  40  cm.  rneist  geringer.  Breite  der  reifsteu  Proglottiden  bis 
8  mm.  Die  Strobila  ist  oft  durch  Liingsfalten  gestreift.  Wohntliier:  Lepus  timid  us. 

This  diagnosis  is  hardly  detailed  enough  to  meet  the  requirements  of 
the  present  day,  but  in  the  anatomical  description  Eiehm  gives  data 
which  supply  what  is  lacking  in  the  passage  just  quoted.  Many  of  the 
points  he  mentions  can  hardly  be  looked  upon  as  specific  characters, 
but  should,  I  believe,  be  attributed  to  influences  of  technique,  individ- 
ual variation,  and  possibly  the  specific  influences  of  environment, 
namely,  the  host — a  subject  to  which  the  helminthology  of  the  future 
must  certainly  give  its  most  serious  and  careful  consideration.  The 
characters  which  appeal  to  me  most  in  Eiehm's  description  may  be 
briefly  summarized  as  follows : 

Male  organs:  The  testicles  form  a  continuous  band  across  the  median  field  in  the 
distal  portion  of  the  segment,  and  in  Plate  VI,  fig.  4,  they  extend  laterally  close  to  the 

JEntozooruui  sive  Vermium  intestinalium  Historia  naturalis,  II,  Pt.  2,  Arnstelse- 
dami,  pp.  82-84. 

2Icones  Helminthum,  pi.  xiv,  figs.  5-6. 
3Systema  Helminthum,  I,  1850,  p.  498. 
4Loc.  cit.,  p.  575. 


NO.  1105.  PROCEEDINGS  OF  THE  NA TIONA L  MVSE  UM.  187 

longitudinal  canals;  the  cirrus  pouch  is  deserving  of  special  notice.  In  the  majority 
of  the  known  Tseniidte  the  cirrus  pouch  is  pyriform  and  rarely  extends  median  of 
the  ventral  canal,  but  in  C.  pectin ata  it  is  a  long  narrow  structure  reminding  one 
of  the  noxzle  of  a  hose;  Riehm  gives  its  average  length  as  more  than  1  mm.,  and  on 
Plate  VI,  fig.  4,  shows  that  it  is  more  than  twice  as  long  as  the  distance  between  the 
genital  pore  and  the  longitudinal  canal.  This  extreme  length  of  the  pouch  is  a 
character  of  great  importance.  Female  organs:  The  vagina,  according  to  Riehm,  is 
about  as  large  as  the  cirrus  pouch.  At  a  point  corresponding  to  the  proximal  end 
of  the  cirrus  pouch  it  becomes  suddenly  very  thin,  and  leads  to  a  second  swelling, 
the  receptaculum  seminis.  The  female  glands  correspond  in  all  essential  characters 
to  those  of  the  genus  Moniezia,  except  that  the  ovary  is  described  as  composed  of 
two  quite  distinct  halves.  Their  position,  some  distance  from  the  longitudinal 
canals,  is  striking.  The  uterus  is  said  to  be  similar  to  that  of  C.  ctcnoides,  namely 
"a  common  uterus  for  both  sides,  which  extends  the  entire  breadth  of  the  segrnen, 
and  is  constricted  only  in  the  middle,  so  that  in  the  gravid  segments  the  lateral 
portion  appears  swollen  by  the  ova  in  comparison  with  the  rather  thin  median 
portion.  Its  volume  is  also  considerably  increased  laterally  [namely,  laterally  to 
the  uterus,  longitudinal  in  reference  to  the  worm]  by  apparently  uubranched  tubes 
which  extend  anteriorly  and  posteriorly.  As  a  matter  of  fact,  however,  these  tubes 
resolve  themselves  as  the  optical  sections  of  a  corresponding  number  of  circular 
wideniugs  of  the  uterus."  Excretory  system:  The  dorsal  canals  become  obliterated 
some  distance  from  the  head.  The  transverse  canals  are  connected  with  one  another, 
not  only  by  the  ventral  canals,  but  also  by  numerous  smaller  longitudinal  canals. 
Topographically,  Riehm  figures  the  genital  canals  as  dorsal  of  the  nerve  and  longi- 
tudinal (ventral)  canal.  The  parasite  is  said  to  occur  only  in  the  fall  and  first  half 
of  the  winter,  and  only  in  hares.  It  was  especially  common  around  the  Roblinger 
See,  but  rare  on  the  higher  plateaux  of  Saxony. 

Blanchard1  in  1891  states  that  he  found  this  species  in  several  hares 
of  unknown  origin.  He  has  never  found  it  in  hares  in  the  central  part 
of  France  or  around  Paris,  but  found  four  specimens  at  Briancon  in 
Lepus  variabilis,  killed  at  a  height  of  1,500  meters.  His  description, 
based  upon  these  specimens,  may  be  summarized  as  follows: 

The  largest  specimen  was  18  cm.  long;  maximum  breadth,  7  to  10  mm. ;  head  315  to 
340  /it  broad ;  neck,  285  to  325  jti  broad ;  in  contracted  specimens  the  neck  may  measure 
1  mm.  broad  at  the  first  segment;  suckers  elliptical,  142  //  long  by  135  JLI  broad; 
opening,  80 // long  by  53 // broad ;  mature  segments,  7  to  10mm.  broad  by  1.1  to  2mm. 
long;  penis  smooth,  40  to  45  f.i  in  diameter,  extrudes  175  to  200  u  from  pore;  eggs 
generally  polygonal  from  reciprocal  pressure,  but  become  elliptical  or  subspherical 
when  pressure  is  removed;  80  to  90  JJL  by  about  75  /.i ;  outer  membrane  1  to  2  u  thick; 
diameter  of  bulb  of  pyriform  body  25  to  30  JLI  ;  length  40  to  50  JLI  ;  horns  terminate  in 
a  long  filament;  hooks  of  oncosphere  8  ju. 

Railliet2  in  1893  places  this  species  in  his  genus  Ctenotwnia. 

Through  the  kindness  of  Dr.  Brandes,  I  have  obtained  one  of  Riehrn's 
original  specimens  of  Dipylidium  pcctinatum  for  comparison  with  the 
American  forms,  and  Blanchard  has  placed  his  forms  from  L.  variabilis 
at  my  disposal.  In  my  private  collection  I  find  several  specimens  ot 
tapeworms  from  Lepus  tlmidus  which  I  collected  in  Leipzig  in  1890,  and 
which  agree  perfectly  with  Rielmrs  form.  With  this  material  at  hand, 
together  with  one  specimen  from  von  Linstow  and  several  from  Moniez. 


'Mem.  Soc.  zool.  France,  IV,  pp.  445,  452,  457-460,  figs.  26-30. 
2Traite  Zool.  uied.  et  agric.,  I,  pp.  278-279. 


188  TAPEWORMS  OF  HAKES  AXD  RABBITS— STILES. 


I  can  add  a  few  points  of  importance  to  Riehm's  diagnosis  and  lay 
greater  stress  upon  other  points  which  1  think  should  be  brought  out 
more  emphatically. 

The  anlagen  of  the  female  glands  and  genital  canals  appear  in  the 
earliest  segments,  while  the  testicles  begin  about  7  mm.  back  of  the 
head.  Male  organs:  The  testicles  are  confined  entirely  to  that  portion 
of  the  segment  which  lies  distally  of  the  uterus,  and  they  extend  on 
both  sides  laterally  of  the  ovaries  to  the  longitudinal  canals ;  this  latter 
character  is  one  of  considerable  importance  when  we  compare  the  Ameri- 
can and  the  European  forms.  Riehm  has  already  directed  attention  to 
the  loughoselike  cirrus  pouch  and  vagina,  but  I  venture  to  call  particu- 
lar notice  to  their  size  and  form. 

Female  organs:  The  excessive  development  which  the  female  glands 
may  attain  is  worthy  of  note:  Riehin's  statement  that  each  ovary 
was  divided  into  two  halves  I  was  unable  to  confirm.  A  peculiar 
feature  of  the  female  glands  is  that  they  develop  suddenly  and  atrophy 
suddenly.  In  Riehm's  cotype,  for  instance,  the  tubes  of  the  ovary  be- 
come visible  8  mm.  from  the  anterior  end  (head  lost) ;  they  then  develop 
rapidly  to  a  maximum  and  suddenly  disappear,  so  that  22  mm.  from  the 
point  where  the  ovarian  tubes  appear  (or  about  30  mm.  from  the  anterior 
extremity)  the  ovary  can  no  longer  be  seen;  the  testicles  persist  very 
much  longer.  The  first  trace  of  the  uterus  is  seen  about  12  mm.  from  the 
anterior  end  of  specimen  (about  14  mm.  from  anterior  end  of  individual,  if 
we  allow  2  mm.  for  the  lost  head  and  first  segments);  the  uterus  passes 
on  the  ventral  side  of  the  ovary,  and  in  all  segments  which  I  have  exam- 
ined there  is  a  single  uterine  anlage  extending  across  the  segment. 

The  genital  canals  and  uterus  run  dorsally  of  the  nerve  and  ventral 
canal  as  Eiehm  figures  them,  and  on  transverse  sections  the  cirrus  is 
shown  to  lie  dorsally  of  the  vagina  on  both  sides  of  the  segment.  The 
nerve  is  very  close  to  the  lateral  margin. 

Kiehm  gives  no  measurements  for  the  ova;  Blanchard  (1891,  p.  460) 
states  that  the  ova  measure  £0  to  90  /;  by  75  /v,  and  that  the  bulb  of 
the  pyriform  body  measures  25  to  30  IJL  in  diameter.  The  measurements 
in  Riehm's  specimen  are  somewhat  smaller,  the  ova  varying  from  72  to 
84  ^  the  bulb  of  the  pyriform  body  14  to  16  y.  My  own  specimens 
agree  with  Riehm's,  but  many  of  the  ova  do  not  exceed  56  ^  in 
diameter. 

The  following  is  proposed  as  a  revised  specific  diagnosis: 

Diagnosis. — Cittotcenia  pectinata  (Goeze,  1782,  partim,  Riehm,  1881), 
Stiles  &  Hassall,  1896.  Strobila  may  attain  400  mm.  in  length  and  8 
mm.  in  breadth,  anterior  portion  usually  lanceolate.  Head  small,  about 
0.25  mm.  or  less  in  diameter,  and  about  0.125  mm.  thick;  rostelliui  and 
hooks  not  observed;  suckers  small.  Neck  very  short,  segmentation 
beginning  almost  immediately  back  of  the  head;  the  proglottids  rapidly 
become  distinct  and  are  always  much  broader  than  long,  the  length  being 
about  one-seventh  of  the  breadth;  gravid  posterior  segments  measure 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  189 


about  10  mm.  broad  by  1.5  mm.  long.  The  anlagen  of  the  genital  organs 
appear  very  early;  the  female  anlagen  are  found  near  the  longitudinal 
canals  immediately  back  of  the  head;  testicles  appear  about  (>  mm. 
back  of  the  head;  genital  pores  double,  in  posterior  half  of  margin. 
Male  organs:  The  testicles  are  about  O.G4  mm.  iu  diameter;  they  are 
numerous,  about  150  in  number,  confined  to  distal  half  of  segment, 
posterior  to  uterus,  and  extend  across  the  entire  median  field  passing 
the  ovary  on  each  side  to  the  longitudinal  canals;  cirrus  pouch  unusu- 
ally large,  attaining  1  mm.  in  length,  extending  some  distance  median 
of  the  longitudinal  canals.  Female  organs:  Ovary,  shell  gland,  und 
vitellogene  gland  situated  some  distance  median  of  the  ventral  canal, 
about  1  mm.  or  more  from  the  lateral  margin;  a  common  transverse 
uterus  to  both  ovaries;  it  passes  the  ovary  ventrally,  is  generally  larger 
in  its  lateral  portions  than  in  its  median  portion,  and  increases  its 
volume  by  proximal  and  distal  branches.  Excretory  system:  Dorsal 
canal  not  observed;  ventral  canal  median  of  nerve;  transverse  canals 
connected  by  secondary  longitudinal  canals.  Longitudinal  nerves  close 
to  lateral  margin.  Cirrus  pouch,  vagina,  and  uterus  pass  from  median 
field  into  lateral  field  dorsally  of  nerve  and  longitudinal  canals.  Cirrus 
pouch  dorsal  of  vagina  on  both  sides  of  the  segment.  Ova  globular  56  to 
84  //.  in  diameter;  bulb  of  pyriform  body  14  to  1C  //;  horns  long,  curved. 

Hosts. — Common  European  hare  (Lepus  timidus)  by  Goeze,  liiehm, 
von  Linstow,  and  Stiles;  Mountain  hare  (L.  variabilis)  by  E.  Blanch- 
ard. Development  unknown. 

Types. — Original  types  1  Cotypes  of  Eiehm  in  the  collection  of 
Leuckart;  No.  1411,  U.S.N.M.  Typical  specimens  in  collection  of 
Blanchard;  collection  of  Moniez;  collection  of  Stiles  (Xos.  116,  1234, 
1238,  U.S.N.M.). 

Geographical  distribution. — Europe:  Germany  (Saxony  by  Eiehm  and 
Stiles;  ?  by  Goeze);  France  (Biiancon  by  It.  Blanchard;  ?  Lille  by 
Moniez). 

CITTOT^NIA    PERPLEXA  (Stiles,  1895),  Stiles  &  Hassall,  1896. 
(Plate  XVIII,  figs.  1-3.) 

1895,  Ctenotcsnia  perplexa,  STILES,  Veterinary  Magazine,  II,  June,  p.  345.     Aug.  28, 1896. 

1896,  Citlotmiia  perplexa  ( STILES,  1895),  STILES  &  HASSALL,  Veterinary  Magazine,  II I, 

p.  407. 

The  specific  name  perplexa  was  proposed  because  it  was  so  difficult 
to  decide  what  to  do  with  the  form  under  discussion.  Some  half  a 
dozen  specimens  were  collected  by  Hassall  from  Lepus  sylvaticus  in 
Bowie,  Maryland.  They  are  all  contracted  and  measure  up  to  57  mm. 
long  by  10  mm.  broad. 

The  parasite  resembles  C.  pectinata  in  general  form  in  the  early 
appearance  of  the  genital  anlagen  and  in  the  fact  that  the  testicles 
extend  beyond  the  ovaries  to  the  lateral  canals.  It  differs  from  C.  pec- 
tinata radically  in  the  size  of  its  cirrus  pouch  and  vagina,  and  in  the 


190  TAPEWORMS  OF  HAKES  AXD  RABBITS— STILES. 


fact  that  the  testicles  are  almost  absent  from  the  median  portion  of 
the  field,  being  arranged  in  two  triangles. 

C.  perplexa  agrees  with  C.  variabilis  in  the  general  size  of  cirrus 
pouch.  It  differs  from  C.  variabilis  in  the  earlier  appearance  of  the 
genital  anlageii  in  the  position  and  arrangement  of  the  testicles.  The 
following  is  proposed  as  specific  diagnosis : 

Diagnosis.— Cittotcenia  perplexa  (Stiles,  1895),  Stiles  &  Hassall,  1896. 
Strobila  attains  57  mm.  long  by  10  mm.  broad.  Head  unarmed,  small, 
about  0.32  mm.  broad,  not  distinctly  separated  from  body;  rostellum 
iiol  observed;  suckers  0.112  mm.  in  diameter.  Keck  extremely  short, 
segmentation  beginning  almost  immediately  back  of  the  head.  Genital 
pores  double,  and  opposite,  in  about  the  middle  of  the  margin.  Anlagen 
of  female  glands  and  canals  visible  within  0.64  mm.  of  the  anterior 
extremity  of  the  head.  Male  organs:  Cirrus  pouch  similar  to  that  of 
C.  variabilis,  but  smaller,  about  0.  -'88  to  0.32  mm.  long,  extending  to  or 
slightly  beyond  the  lateral  nerves;  testicles  arranged  in  two  groups  in 
each  segment,  one  triangle  being  around  each  ovary  and  extending 
laterally  to  the  longitudinal  canals.  Female  organs :  Agree  essentially 
with  those  of  C.  variabilis,  as  does  the  general  topography;  uterus 
single  or  double. 

Host. — Cottontail  Babbit  (Lcput>  sylvaticus]  by  E assail  in  Bowie, 
Maryland. 

Types. — Bureau  of  Animal  industry,  Cestode  series,  No.  1126,  desig- 
nated as  type,  and  deposited  in  the  United  States  National  Museum. 
Paratypes  in  Bureau  of  Animal  Industry;  Nos.  1110,  1131,1137-1139, 
U.S.N.M.;  collection  of  Stiles;  collection  of  Hassall.  Other  typical 
specimens  will  not  be  distributed  until  more  material  is  obtained. 

CITTOT^ENIA  VARIABILIS  (Stiles,  1895),  Stiles  &  Hassall,  1896. 
(Plate  XIX,  figs.  1-14;  Plate  XX,  figs.  1-5.) 

1892,  "Tcenia  pectinala,  GOEZE,"  1782,  ex  parte  of  CURTICE,  Journ.  Comp.  Med.  Vet. 
Arch.,  XIII,  pp.  232-233. 

1895,  CtenotceniavariabiUs,  STILES, Veterinary  Magazine,  II.  June,  p.  345.    Aug.  28, 1896. 

1896,  Cittotcenia  variabilis  (&TILKS,  1895),  STILES  &  HASSALL,  Veterinary  Magazine, 

III,  p.  407. 

As  stated  in  the  introduction,  helminthology  is  not  so  far  advanced 
that  it  is  possible  for  us  to  determine  what  limits  should  be  given  to 
genera,  subgeuera,  species,  and  subspecies,  and  for  some  time  to  come 
all  classification  into  groups  must  be  looked  upon  as  experimental,  the 
ideas  of  every  author  being  subject  to  change  from  day  to  day  as  new 
facts  in  the  comparative  anatomy  of  cestodes  are  published  or  observed. 
The  American  form  which  I  now  describe  as  Cittotcenia  variabilis  is 
one  of  the  parasites  which  can  equally  well  be  considered  as  a  distinct 
species,  or  as  a  subspecies,  possibly  peculiar  to  given  hosts.  It  is  so 
perfectly  distinct  from  the  European  C.  pectinata  that  no  specialist 
could  fail  to  recognize  the  differences  when  he  has  the  two  forms  side 
by  side  for  comparison.  Yet  it  is  so  closely  allied  to  the  European 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  191 

C.  pectinata  that  the  question  forces  itself  upon  us  whether  these  dif- 
ferences can  not  be  considered  as  subspecific,  possibly  due  to  some 
extent  to  a  difference  of  conditions  found  in  different  hosts  and  in 
different  countries.  I  frankly  admit  that  during  an  examination  of  a 
large  series  of  specimens  1  have  changed  my  mind  a  dozen  times  in 
regard  to  tliis  particular  species.  I  am  now  of  the  decided  opinion, 
however,  that  this  form  should  be  given  specific  rank. 

The  specimens  upon  which  this  description  is  based  were  collected 
at  Bowie,  Maryland,  by  Hassall  from  the  cottontail  (Lepm  Ni/lraticus). 
The  strobila  attains  100  to  180  mm.  in  length  and  10  mm.  in  breadth. 

The  head  is  very  small,  measuring  0.32  to  0.56  mm.  broad;  it  may  or 
may  not  be  sharply  denned  from  the  neck;  no  rostellum  or  hooks  visible; 
the  suckers  are  small,  measuring  0.12  to  0.28  mm.  long  by  0.112  to  0.24 
rnm.  broad;  the  opening  of  the  sucker  is  directed  diagonally  forward. 
The  neck  is  very  short  or  absent,  segmentation  beginning  very  early  and 
the  proglottids  rapidly  becoming  distinct.  The  anterior  portion  varies 
greatly  in  shape,  according  to  contraction.  The  genital  anlagen  appear 
rather  early;  at  about  5  to  10  mm.  from  the  head  two  roundish  bodies 
appear  in  each  segment,  one  on  each  side  of  the  median  field  near  the 
longtndinal  canals;  these  anlagen  soon  change  in  form  to  the  pistol- 
shaped  anlagen  described  for  Moniezia;  at  about  25  mm.  from  the 
head  the  first  testicles  are  visible.  The  genital  pores  are  double  and 
opposite,  and  situated  in  about  the  middle  third  of  the  margin. 

Segments  125  mm.  from  the  head  measure  0.56  mm.  long  by  6.5  mm. 
broad  by  0.48  mm.  thick.  Male  organs:  The  testicles  number  about  65 
to  90;  they  are  confined  to  the  dorsal  half  of  the  segment  distally  of 
the  transverse  uterus,  and  do  not  extend  laterally  of  the  ovaries.  The 
cirrus  pouch  lies  dorsally  of  the  vagina  on  both  sides  of  the  segment; 
it  is  long,  narrow,  and  quite  indistinct,  coloring  in  carmine  much  more 
lightly  than  the  vagina.  Female  organs:  The  vagina  runs  ventrally 
of  the  cirrus;  it  is  long  and  narrow.  From  the  margin  of  the  segment 
for  a  distance  of  about  0.48  mm.  it  is  surrounded  by  a  deeply  coloring- 
layer  of  cells,  and  then  it  is  reduced  to  a  thin  narrow  canal  which  later 
swells  into  the  receptaculum  seminis.  The  ovary,  shell  gland,  and 
vitellogene  gland  resemble  those  of  Moniezia  and  C.  pectinala  and  lie 
1.28  mm.  from  the  lateral  margin.  The  uterus  may  be  double  or  single; 
in  some  segments  a  single  uterine  aulage  extends  across  the  entire  seg- 
ment, running  through  the  ventral  portion  of  the  ovary,  and  passing 
to  the  lateral  fields  dorsally  of  the  longitudinal  canals  and  nerves;  in 
the  majority  of  segments  there  are  two  distinct  uterine  anlagen,  one 
to  each  set  of  female  glands.  The  ventral  canal  runs  about  midway 
between  the  ovary  and  the  lateral  margin;  the  dorsal  canal  lies  dorso- 
median  of  the  ventral  canal  and  is  bound  by  a  heavy  cuticular  lining. 
The  longitudinal  nerve  lies  laterally  of  the  ventral  canal,  and  ventrally 
of  the  genital  ducts,  about  0.64  mm.  from  the  lateral  margin. 

In  the  older  segments  the  topography  described  above  is  preserved, 


192  TAPEWORMS  OF  HARES  AXD  RABBITS- STILES. 


but  gradually  the  uterus  develops  to  such  an  extent  that  it  suppresses 
all  the  genital  glands.  The  cirrus  pouch  becomes  indistinct,  bat  the 
vagina  can  be  traced  in  nearly  all  segments. 

As  the  uterus  increases  in  size  it  gives  rise  to  proximal  and  distal 
branches,  but  the  latter  are  far  less  regular  than  those  of  the  European 
C.  pectinata.  In  the  posterior  segments  it  is  generally  difficult  to  dis- 
tinguish the  two  uteri.  The  ova  measure  CO  to  04  //  in  diameter:  the 
bulb  of  the  pyriform  body,  12  to  16  /,/.  This  is  the  species  (var.  angusta] 
upon  which  my  paper.  "A  double-pored  cestode  with  occasional  single 
pores77  was  based,  and  since  writing  that  note  another  case  of  the  same 
variation  was  noticed.  On  account  of  the  numerous  variations  in 
the  position  of  the  genital  pores,  uterus,  etc.,  noticed  in  the  specimens 
studied,  I  proposed  to  name  the  worm  Ctenotcenia  rariabilis. 

Larval  stage. — The  young  specimens  described  on  page  201  were  col- 
lected in  the  same  locality  as  the  species  here  described,  and  from  the 
same  host;  and  it  seems  probable  that  the  unarmed  young  mentioned 
on  page  201  are  the  young  of  either  C.  rariabilis  or  C.  perplexa. 

Diagnosis. — Cittotcenia  variabilis  (Stiles,  1895),  Stiles  &  Hassall,  1896. 
(American  representative  of  C.  pectinata.)  Strobila  attains  100  to 
180  mm.  in  length  and  10  mm.  in  breadth.  Head  small,  about  0.3  to 
0.6  mm.  in  breadth ;  it  may  or  may  not  be  denned  from  the  neck  accord- 
ing to  contraction.  Neck  very  short,  segmentation  beginning  almost 
immediately  back  of  the  head;  segments  always  much  broader  than 
long.  Genital  anlagen  appear  very  early,  about  5  to  10  mm.  from  the 
head.  Genital  pores  double,  in  about  the  middle  third  of  the  lateral 
margin.  Male  organs:  Testicles  about  60  to  100  in  each  segment,  con- 
fined between  the  ovaries  to.  the  dorsal  portion  of  the  distal  half  of  the 
median  field;  cirrus  pouch  about  0.4  mm.  long,  narrow  and  very  indis- 
tinct, lying  dorsally  of  the  vagina  on  both  sides  of  the  segment.  Female 
organs:  Ovary,  shell  gland,  and  vitellogene  gland  resemble  the  corre- 
sponding organs  of  Moniezia;  they  lie  median  of  longitudinal  canals, 
about  1.3  mm.  from  the  lateral  margin;  the  vagina  is  rather  distinct, 
lies  ventrally  of  the  cirrus  pouch,  and  for  about  0.5  mm.  from  the  pore 
it  is  surrounded  by  deeply  staining  cells;  uterus  may  be  double  or 
single  (in  the  same  specimen)  and  may  produce  proximal  and  distal 
branches;  ova  60  to  64  /*  in  diameter;  bulb  of  pyriform  body  12  to  16  //. 
Longitudinal  nerve  about  0.6  mrn.  from  the  lateral  margin;  ventral 
canal  large  with  thin  lining;  dorsal  canal  much  smaller  than  ventral 
canal,  dorso-median  of  ventral  canal,  with  thick  lining;  transverse 
canals  connect  ventral  canals  [no  injections  made  for  secondary  longi- 
tudinal canals].  Genital  ducts  and  uterus  pass  from  median  to  lateral 
field  on  the  dorsal  side  of  the  longitudinal  canals  and  nerves. 

Host. — Cottontail  Babbit  (Lepus  syh'aticus),  by  Hassall;  Marsh  Hare 
(L.  palustris),  by  Mills. 

Types. — Bureau  of  Animal  Industry,  Cestode  series  No.  117  desig- 
nated as  type  and  deposited  in  the  United  States  National  Museum. 
Paratypes  distributed  as  follows:  Collection  of  Bureau  of  Animal 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  193 

Industry-  collection  of  Leidy  (University  of  Pennsylvania);  collection 
of  Harvard  Museum  of  Comparative  Zoology;  collection  of  Ward;  col- 
lection of  Stiles ;  collection  of  Hassall.  Europe :  Berlin  Museum ;  collec- 
tion of  Leuckart;  collection  of  Max  Braun ;  Halle  Zoological  Institute; 
Vienna  Museum;  collection  of  Stossich;  collection  of  Parona;  collec- 
tion of  Monticelli;  collection  of  Zschokke;  collection  of  R.  Blanchard; 
collection  of  Eailliet;  collection  of  Neumann;  collection  of  Moniez; 
British  (South  Kensington)  Museum.  Asia  :  Imperial  University 
of  Japan,  Tokyo.  Technique  of  types :  Corrosive  sublimate,  acetic  acid, 
acid  carmine. 

Unarmed  young  stages  deposited  in  collection  of  Bureau  of  Aiiimal  Industry;  col- 
lection of  U.S.N.M. ;  collection  of  Stiles;  collection  of  Hassall;  collection  of  Berlin 
Museum;  collection  of  Leuckart;  collection  of  R.  Blanchard. 

Geographical  distribution. — Maryland  (by  Hassall),  Florida  (by  Mills), 
Long  Island  (by  Peters),  ?  Puget  Sound  (collection  of  Leidy). 

Varieties. — To  classify  the  forms  at  my  disposal,  I  am  compelled  to 
recognize  three  varieties : 

(a)  C.  variabilis.    B.  A.  I.  Cestode  series  No.  117,  type  of  the  species 
is  designated  as  type  of  this  variety.     The  posterior  liaps  of  the  seg- 
ments are  nearly  straight;  genital  pore  in  about  the  middle  of  the  lat- 
eral margin.     Habitat:  Lepus  sylvaticus. 

(b)  C.  variabilis  angusta.    B.  A.  I.  Cestode  series  No.  1119,  designated 
as  type  and  deposited  in  the  United  States  National  Museum.    This 
variety  is  only  about  2  mm.  broad,  the  posterior  flap  is  straight  and  does 
not  overlap  prominently,  the  genital  pore  is  generally  in  the  posterior 
half  of  the  lateral  margin.     Pores  occasionally  single. 

(c)  C.  variabilis  imbricata.     No.  1246,  U.S.N.M.,  borrowed  by  B.  A.  I.; 
B.  A.  I.  Cestode  series  No.  1246,  designated  as  type  and  returned  to 
United  States  National  Museum.    At  first  sight  it  seems  almost  like 
splitting  hairs  to  create  a  variety  for  these  specimens  the  technique  of 
which  was  different  from  that  of  C.  variabilis.    The  worms  were  col- 
lected by  Kobert  Mills,  an  enthusiastic  collector  at  Chuluota,  Florida, 
and  kindly  presented  to  the  United  States  National  Museum.    They 
were  placed  in  95  per  cent  alcohol,  accordingly  they  are  somewhat  con- 
tracted.    The  posterior  border  of  the  segments  is  lobed,  a  character 
which  is  quite  constant,  and  overlaps  the  next  following  segment  about 
one-third  of  its  length  so  that  the  segments  have  a  general  campulate 
appearance. 

Subfamily    DliPYILIDIIlST^E,    Railliet,    1896. 

1850,  Section  Ehynchotcenia,1  DIESING,  Sy sterna  Helminthum,  I,  p.  497. 

1858,  Subf.  Malacolepidota  (Soft-shell Tapeworms),  WEINLAND,  Human Cestoides,  p.  52. 

1863,  Cystoidew,  R.  Leuckart,  Die  menschlichen  Parasiten,  I,  p.  389. 

1864,  Subg.  Rhynchotwnia,1  DIESING,  Sitzber.  Akad.  Wiss.  Wien,  XLIX,  1,  p.  365. 

iKhynchotwnia,  Diesing,  1850,  a  "  section"  of  Toenia,  may  be  interpreted  as  a  sub- 
genus;  by  the  law  of  priority,  however,  it  falls  as  a  synonym  of  Fimbriaria,  Frolich, 
1802,  taking  of  course  the  same  species  as  type,  i.  e.,  Tceuia  malleus,  Goeze,  1782.  In 
order  to  meet  objection  to  this  ruling,  should  objection  arise,  I  here  definitely  pro- 
pose  Tcenia  malleus,  Goeze,  as  type  of  the  subgenus. 

Proc.  N.  M.  vol.  xix 13 


194  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 

1884,  Subg.  Microtcenia,1  SEDGWICK,  in  GLAUS  &  SEDGWICK,  Elementary  Text-book 

of  Zoology,  I,  p.  336. 

1886,  Cystoidei,  K.  LEUCKART,  Die  Parasiten  des  Menschen,  2d  ed.,  I,  p.  825. 
1886,  Group  Cystoidotcenice,  RAILLIET,  filaments  de  Zool.  m6d.  et  agric.,  p.  253. 
1896,  DipyUduncc,  EAILLIET,  Recueil  de  Me~d.  vdt.,  8  ser.,  Ill,  5,  p.  159.— STILES,  Bull. 

12,  Bureau  of  Animal  Industry,  U.  S.  Dept.  Agric.,  1896,  p.  28. 

Diagnosis. — Taeniidse  with  rostellum  which  is  generally  armed  ;  geni- 
tal pores  lateral  (marginal),  single  or  double;  eggs  with  thin  transparent 
shells,  frequently  arranged  in  egg  sacs,  in  some  cases  scattered  through 
the  segments  ;  larval  stage  a  cysticercoid ;  adults  in  birds  and  mam- 
mals. 

Type. — Dipylidium,  E.  Leuckart,  1863. 

Genus  DAVAINEA,  R.  Blanchard  &  A.  Railliet,  1891. 

1891,  Davainea,  R.  BLANCHARD  &  RAILLIET,  Me~m.  Soc.  zool.  France,  IV,  p.  428. 

Type,  Tcenia  proglottina,  Davaine,  1860. 
?1893,  Chapmania,  MONTICELLI,  Naturalista  Siciliano,  XII  (7-8-9),  pp.  16-19.     Type, 

Tcenia  ar</e»/ina,-Zschokke,  1888. 

Diagnosis. — Dipylidiinre  of  small  or  medium  size.  Head  surmounted 
by  a  rostellum  or  hollowed  by  a  depression,  but  armed  in  either  case 
with  a  double  row  of  numerous  small  hooks  of  special  form  (prong  and 
dorsal  root  short,  ventral  root  very  long,  giving  to  the  hooks  the  form 
of  a  hammer).  Suckers  bordered  with  several  rows  of  small  hooks 
which  may  be  instable  or  persistent.  Genital  pores  unilateral  or  irreg- 
ularly alternate;  in  the  former  case  the  ova  are  generally  arranged  in 
egg  capsules;  in  some  species  the  ovary  develops  into  uterus;  eggs  may 
also  be  isolated,  scattered  through  the  parenchyma. 

Type. — Davainea  proglottina  (Davaine,  1860),  E.  Blanchard,  1893. 

Habitat. — Adults  in  intestine  of  birds  and  mammals;  cysticercoids, 
generally  in  arthropods  and  mollusks. 

This  genus  is  found  chiefly  in  birds,  but  two  species  have  been 
recorded  in  mammals,  besides  the  two  species  here  given  for  Lepus. 
The  diagnoses  of  the  two  other  forms,  which  are  introduced  for  com- 
parison, read  as  follows  : 

DAVAINEA  MADAGASCARIENSIS  (Davaine,  1870),  R.  Blanchard,  1891. 

1870,  Tcenia  madagascariensis,  DAA^AINE,  M6m.  Soc.  Biol.  Paris,  5  ser.,  I  (1869),  1870, 

pp.  233-240,  pi.  I 
1891,  Davainea  madagascariensis  (DAVAINE,  1870),  R.  BLANCHARD,  Me"m.  Soc.  zool. 

France,  IV,  pp.  429,  438. 

Diagnosis. — Davainea  madagascariensis  (Davaine,  1870),  E.  Blanch- 
ard, 1891.  Strobila  attains  250  to  300  mm.  in  length,  and  is  formed  of 
500  to  600  trapezoidal  segments  which  are  broader  than  long.  Head 

:The  subgenus  Microtcenia  of  the  genus  Tcenia  contains  the  species  Tcenia  cncu- 
merina,  T.  elliptica,  T.  nana,  and  T.  flavo-punctata,  and  is  thus  antedated  by  the  genera 
Dipylidium,  Leuckart,  1863,  Diplacantlms,  Weinland,  1858  (nee  L.  Agassiz),  and 
Hymenolepis,  Weinland,  1858,  but  the  type  has  never  been  proposed.  In  order  to 
derinitely  dispose  of  the  genus  I  propose  Tcenia  cucumerina  (=  Tcenia  canina}  as  type, 
thus  making  Microtcenia  a  synonym  of  Dipylidium. 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM. 


195 


provided  with  rather  a  large  rostellum  armed  with  a  double  crown  of 
about  90  hooks  18  /.i  long.  Suckers  round  and  quite  large  5  hooks  on 
suckers  not  observed.  Genital  pores  unilateral.  A  single  egg  in  each 
egg  sac. 

Hos L— Man  (Homo  sapiens).  Found  in  Mayotte,  Mauritius,  and 
Bangkok. 

DAVAINEA   CONTORTA,  Zschokke,  1895. 
(Plate  XXII,  fig.  2.) 

1895,  Davainea    contorta,    ZSCHOKKE,  Centralbl.    f.     Bakter.  u.    Parasitenk.,  Pt.  I, 
XVII  (18-19),  pp.  634-645,  figs.  1-4.     21  May. 

Diagnosis. — Davainea  contorta,  Zschokke,  1895.  Strobila  attains  40 
to  80  mm.  in  length  by  0.75  mm.  in  breadth,  and  contains  400  to  800 
segments,  all  of  which  are  broader  than  long 5  margin  of  strobila  ser- 
rate. Scolex  small,  prismatic;  rostellum  large,  armed  with  (?  a  single 
row  of)  numerous  minute  hooklets;  suckers  large,  armed  with  8  to  10 
rows  of  minute  hooklets.  Genital  organs  differentiate  in  fortieth  to 
sixtieth  segment.  Genital  pores  unilateral,  in  middle  of  lateral  mar- 
gin. Male  organs:  Cirrus  pouch  one-fourth  to  one  third  as  Jong  as  the 
segment  is  broad;  testicles  two,  large,  in  dorsal  portion  of  median  field 
on  aporose  side  of  median  line.  Female  organs :  Ovary  bilobed ;  gravid 
segments  filled  with  large  egg  capsules,  each  with  a  single  egg  pos- 
sessing two  shells. 

•  Host. — Common  Indian  Pangolin  (Manis  pentadactyla).    Type  with 
Zschokke. 

DAVAINEA  RETRACTILIS,  Stiles,  1895. 

(Plate  XXI,  figs.  1-6;  Plate  XXII,  fig.  1.) 
1895,  Davainea  retractilis,  STILES, Veterinary  Magazine,  II,  June,  p.  343.     Aug.  28, 1895. 

Of  this  species,  I  have  seven  strobile,  collected  March  10, 1891,  in 
Nevada,  by  Dr.  Fisher,  of  the  Division  of  Mammalogy  and  Ornithology, 
United  States  Department  of  Agriculture.  The  original  label  reads: 
"Biological  Explorations,  U.  S.  Dept.  Agr.,  Death  Valley  Expedition, 
Ash  Meadows,  Nevada,  Mch.  10,  1891.  A.  K.  Fisher.  Nye  Co.  Tape- 
worms from  Cottontail,  No.  362."  A  reference  to  the  specimen  in  the 
Division  of  Mammalogy  and  Ornithology  shows  that  the  host  is  Lepus 
arizonce. 

Measurements  of  Davainea  retractilis. 


B.  A.  I.  Cestode 
series  (No.). 

Length. 

Breadth. 

Condition. 

1168 

mm. 
35 

mm. 
3 

Fragment. 

1187          

65 

3 

With  head. 

1188 

98 

3 

Fragment. 

1189                

80 

2.75 

With  head  ;  type. 

1190 

j           68 

2.75 

Do. 

1191            

74 

2.5 

Fragment. 

1192 

105 

3 

With  head. 

196  TAPEWORMS  OF  HARES  AND  RABBITS— STILES. 


All  of  the  specimens  are  so  contracted  that  a  study  of  the  general 
anatomy  in  detail  is  out  of  the  question;  enough  can  be  given,  however, 
to  clearly  define  the  species  from  other  members  of  this  genus.  Tak- 
ing the  type  specimen  as  basis  for  description,  I  find  the  following 
characters: 

The  head  measures  0.480  mm.  broad  by  0.32  mm.  long.  The  rostellnm 
measures  80  //  in  its  trau verse  and  48  JJL  in  its  longitudinal  diameter; 
its  equator  is  provided  with  a  crown — on  other  preparations  shown  to 
be  double — of  minute  booklets  12  p  long.  It  is  impossible  to  count  the 
hooks,  but  there  are  about  as  many  present  as  in  I),  salmoni — i.  e.,  about 
90  to  120.  The  suckers  are  oblong  and  measure  0.2  to  0.224  inm.  by  0.14 
to  0.16  mm.;  they  are  armed  with  numerous  booklets  arranged  in  rows. 
An  estimate  of  their  number  is  even  more  difficult  than  in  the  case  of 
Davainea  salmoni  (see  p.  198).  There  are  about  75  diagonal  rows  of 
roots;  from  5  to  20  roots  have  been  counted  in  different  rows.  Five 
hundred  to  seven  hundred  hooks  would  probably  not  be  a  high  esti- 
mate for  each  sucker.  The  suckers  appear  at  first  sight  with  low  pow- 
ers, to  open  at  right  angles  to  the  longitudinal  surface  of  the  worm, 
but  a  careful  study  shows  that  the  entire  muscular  bulb,  together  with 
its  numerous  hooks,  is  retracted  into  a  sac-like  structure  which  opens 
directly  forward — namely,  at  right  angles  to  a  transverse  section;  in 
other  words,  the  entire  sucker  is  inverted  in  the  parenchyma  of  the 
head ;  the  direction  of  the  opening  of  this  invagination  varies  slightly 
in  different  specimens.  A  circular  canal  9  yu  in  diameter  surrounds  the 
rostellurn;  a  longitudinal  canal  runs  in  the  dorsal  and  another  in  the 
ventral  median  line,  which  in  all  probability  are  connected  with  the  cir- 
cular canal,  although  this  could  not  be  absolutely  demonstrated. 

The  contraction  was  such  that  the  neck  and  the  commencement  of 
stabilization  could  not  be  judged  satisfactorily,  although  the  neck  is 
apparently  short,  segmentation  beginning  near  the  head.  As  nearly  as 
could  be  estimated,  the  strobila  contains  about  950  segments;  in  this 
estimate  about  300  segments  are  allowed  for  the  first  10  mm.  of  the  worm. 
In  the  anterior  portion,  the  four  longitudinal  canals  are  very  distinct 
and  laterally  the  longitudinal  nerve  is  indistinctly  visible;  the  larger 
(evidently  ventral)  canal  is  lateral  of  the  smaller  (dorsal)  canal.  About 
1.12  mm.  from  the  head  a  small  deeply- staining  line  of  cells  develops  in 
the  median  line  of  the  segments,  running  parallel  to  the  anterior  mar- 
gin; this  gradually  increases  in  length  as  the  segments  are  followed 
distally:  this  is  interpreted  as  the  anlage  of  the  genital  canals  and 
probably  also  of  the  female  glands.  Very  shortly  after  its  appearance 
two  groups  of  testicles  are  noticed,  one  group  each  side  of  the  median 
line.  The  cirrus  pouch  and  vagina  appear  in  the  lateral  field,  all  of 
the  genital  pores  being  unilateral  in  the  anterior  half  of  the  segment. 
The  pouch  is  small,  measuring  0.12  mm.  long  by  60  ^  broad;  no  clearly 
defined  vesicula  seminalis  could  be  seen,  but  the  coils  made  by  the 
retracted  cirrus  appear  to  occupy  the  entire  pouch;  the  vagina  is 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  197 

immediately  distal  of  the  pouch.  The  changes  in  the  genital  system  must 
be  left  for  some  one  to  study  AV!IO  can  preserve  fresh  material  for  this  pur- 
pose. Gradually  the  entire  median  field  becomes  filled  with  eggs; 
while  several  eggs  are  contained  in  each  capsule  in  I),  salmoni,  the  ar- 
rangement in  7).  retractilis,  so  far  as  could  be  judged  from  the  poor 
material,  agrees  more  closely  with  that  described  for  I),  contorta  and  D. 
madagaxcariensis,  each  capsule  containing  but  one  egg.  The  ova  are 
about  80  J.L  in  diameter  and  possess  two  (?)  surrounding  membranes. 

The  segments  are  all  broader  than  long,  variations  being  found 
between  0.56  mm.  broad  by  4  j.i  long  (anterior  segments)  and  3  mm.  broad 
by  0.24  mm.  long.  The  distal  segments  are  slightly  narrower,  but  there 
is  not  the  sudden  and  distinct  change  seen  in  D.  salmoni.  As  specific 
name  I  have  proposed  Davainea  rctractilis,  referring  to  the  retracted 
condition  of  the  suckers  found  upon  all  the  heads  examined. 

The  material  at  hand  does  not  warrant  further  description,  but  from 
the  data  given  the  following  diagnosis  is  proposed: 

Diagnosis. — Davainea  retractilis,  Stiles,  1895.  Strobila  105  mm.  or 
more  long  by  3  mm.  broad,  with  about  1,000  segments,  all  of  which  are 
much  broader  than  long;  segments  vary  from  0.56  mm.  broad  by  4  /,*  long 
(anterior  segments)  to  3  mm.  broad  by  0.24  mm.  long  (gravid  segments). 
Head  measures  0.37  to  O.G8  mm.  broad  by  0.36  to  0.43  mm.  long.  Ee- 
tractile  rostellum  80  //  by  48  /./,  armed  with  a  double  row  of  minute 
booklets  12  /,/  long,  about  40  to  60  books  in  each  row.  Suckers  meas- 
ure 0.2  by  0.14  mm.,  and  may  be  completely  retracted  into  the  bead; 
they  are  armed  with  numerous  booklets  arranged  in  rows;  the  roots 
arrange  themselves  regularly  in  rows,  about  75  diagonal  rows  coming 
to  each  sucker,  and  5  to  20  roots  in  each  row ;  about  500  to  700  hooks 
are  estimated  for  each  sucker.  Neck  is  short  or  absent,  strobilization 
beginning  almost  immediately  back  of  the  head.  Longitudinal  canals 
at  first  very  distinct,  ventral  canal  lateral  of  dorsal  canal.  Genital 
anlage  appears  about  1  mm.  from  the  head,  and  is  at  first  median;  tes- 
ticles arranged  in  two  groups,  one  each  side  of  the  median  line.  Gen- 
ital pores  unilateral,  cirrus  pouch  small,  0.12  mm.  long  by  60  /*  broad; 
vagina  immediately  distal  to  pouch.  Eggs  80  //  in  diameter,  inner  shell 
40  /^;  a  single  egg  in  each  egg  capsule. 

Host. — One  of  the  cottontail  rabbits  (Lepus  arizoncc)  by  A.  K.  Fisher; 
development  not  known. 

Types. — Diagnosis  based  upon  four  strobilse  with  beads,  and  several 
fragments;  Bureau  of  Animal  Industry  Cestode  series  No.  1189  is  des- 
ignated as  type  and  deposited  in  the  United  States  National  Museum. 
Paratypes  distributed  as  follows :  Collection  of  Bureau  of  Animal  In- 
dustry; Berlin  Museum;  Vienna  Museum;  collection  of  E.  Blancbard. 
Fragments  to  British  Museum;  collection  of  Parona  (Genoa);  Tokyo 
University;  collection  of  Zscbokke  (Bale).  All  poorly  preserved;  tech- 
nique, alcbobol  and  hseinatoxylin  or  acid  carmine. 

Geographical  distribution. — Nevada,  by  A.  K.  Fisher. 


198  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 

DAVAINEA  SALMONI,  Stiles,  1895. 

(Plate  XXII,  figs.   3-4;  Plate  XXIII,  figs.  1-9;  Plate  XXIV,  figs.  1-2 ;   Plate  XXV, 

figs.  1-11.) 

1887,  "  Tcenia  pectinata,"  GOEZE,  1782,  of  CURTICE  exparte,  Science  [N.  Y.],  March  23. 
1895,  Davainea  salmoni,  STILES,  Veterinary  Magazine,  II,  June,  p.  343.     Aug.  28, 1895. 

The  length  of  type  specimen,  mounted  (Bureau  of  Animal  Industry 
Cestode  series  No.  1196),  is  86  mm.;  the  broadest  segments  attain  3  mm. 
in  width.  Total  number  of  segments  about  450  (the  exact  number  could 
not  be  ascertained  because  of  contraction  at  certain  points),  of  which 
about  230  belong  to  the  first  third  of  the  worm.  Head  0.736  mm.  broad 
by  0.496  long.  Retracted  rostellum  0.144  mm.  in  diameter,  provided 
with  a  double  crown  of  minute  hooks,  of  characteristic  Davainea  form, 
20  j.i  long,  about  60  hooks  in  each  row.  The  suckers  are  large  and 
prominent  and  armed  with  rows  of  closely  set  hooks,  the  number  of 
which  it  is  utterly  impossible  to  count;  they  may  be  estimated  at  about 
750;  the  size  of  these  hooks  varies,  the  prong  of  the  longest  reaching 
10  jj.  The  neck  is  thin  (0.480  mm.  broad)  and  short,  segmentation 
beginning  about  0.8  mm.  back  of  the  head.  The  segments  vary  extremely 
according  to  contraction,  most  of  them  being  broad  and  short,  but 
others  being  infundibuliform  and  nearly  as  long  as  broad.  Within  the 
proximal  third  of  the  strobila  variations  may  be  found  between  seg- 
ments 0.512  mm.  broad  by  32  fj.  long  and  0.7  mm.  broad  by  0.528  mm. 
long. 

.The  anlage  of  the  genital  organs  first  appears  in  about  the  one 
hundred  and  seventieth  segment,  as  a  darkly  staining  body,  one  end 
of  which  is  in  the  median  line,  the  other  pointed  toward  the  pore  side 
of  the  segment.  As  the  segments  are  followed  distally  this  body 
assumes  the  characteristic  pistol  shape  found  in  the  genital  anlagen  of 
so  many  tapeworms  (Moniezia,  etc.);  the  muzzle  of  the  barrel  rapidly 
reaching  the  lateral  margin  at  the  genital  pore;  the  barrel  divides  into 
two  parallel  canals,  the  vas  deferens  proximally,  the  vagina  distally, 
while  the  handle  of  the  pistol  grows  distally  in  the  median  line  to  form 
the  female  glands  in  approximately  the  middle  of  the  segment.  The 
genital  pores  are  irregularly  alternate,  and  vary  in  position  from  the 
middle  to  near  the  anterior  edge  of  the  margin.  The  testicles  appear 
at  about  the  two  hundred  and  thirtieth  segment,  and  occupy  almost 
the  entire  median  field.  The  condition  of  the  material  does  not  warrant 
a  description  of  the  changes  the  female  anlage  undergoes,  further  than 
to  state  that  the  glandular  portion  divides  into  two  (or  probably  three) 
bodies.  The  anterior  body  will  by  analogy  represent  the  ovary,  the 
distal  body  the  vitellogene  gland;  the  third  (very  indistinct)  body 
would  probably  represent  the  shell  gland.  Canals  could  be  seen 
between  these  glands,  but  could  not  be  analyzed.  The  development 
of  the  uterus  could  not  be  followed,  but  after  about  the  three  hundred 
and  fortieth  segment  groups  of  eggs  gradually  appear,  suppressing 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSETM.  199 


all  of  the  genital  glands.  The  cirrus  pouch  and  vagina  could  not  be 
studied  in  detail;  the  former  is  very  small  and  muscular,  and  measures 
0.14  mm.  long  by  44  /t  broad.  Excretory  and  nervous  systems  could 
not  be  analyzed. 

These  are  all  the  details  which  can  be  given  for  the  type  specimen, 
but  from  some  of  the  other  material  the  following  could  be  observed: 

The  vagina  is  distal  of  the  cirrus  pouch.  The  muscular  layer  of  the 
cirrus  pouch  measures  up  to  20  u  thick.  The  cirrus  when  retracted 
forms  several  coils  in  the  pouch,  but  no  vesicula  seminalis  could  be 
distinguished;  in  fact  there  is  no  room  for  one  within  the  pouch  when 
the  cirrus  is  retracted.  The  egg  capsules  fill  the  entire  median 
field  of  the  segment,  and  occasionally  extend  into  the  lateral  fields; 
there  are  about  160  visible  upon  one  face  of  a  segment;  they  are 
globular  in  form,  but  assume  various  shapes  by  reciprocal  pressure; 
they  measure  0.112  to  0.128  mm.  in  diameter  and  contain  3  to  15  ova; 
the  ova  measure  20  to  24  /*  in  diameter. 

The  posterior  10  to  20  segments  decrease  in  breadth  and  increase  in 
length  very  rapidly,  in  some  cases  measuring  1.8  mm.  broad  by  1.4  mm. 
long. 

In  some  specimens  the  pores  show  a  remarkable  tendency  to 
unilaterality;  in  fact,  in  two  specimens  which  Dr.  Norgaard  has  col- 
lected since  this  article  was  written  all  of  the  pores  are  on  the  same 
side  of  the  worm.  The  position  of  the  genital  pore  is  evidently  a  very 
uncertain  character  in  the  genus  Davainea. 

Further  details  are  not  warranted  by  the  material  at  hand. 
*  As  a  name  for  this  parasite,  I  proposed  Davainea  salmoni,  dedicating 
the  species  to  my  friend  and  chief,  Dr.  Daniel  E.  Salmon,  Chief  of  the 
Bureau  of  Animal  Industry,  United  States  Department  of  Agriculture, 
to  whose  broad  policy  of  administration  1  am  indebted  for  the  opportu- 
nities of  carrying  on  my  work  in  helminthology. 

The  larval  stage. — In  December,  1887,  Cooper  Curtice  examined  a 
rabbit  (Lepus  sylvaticus)  in  which  he  found  a  numb.er  of  tapeworms 
in  various  stages  of  development.  He  made  an  extremely  important 
observation  on  the  younger  specimens,  which,  unfortunately,  he  never 
published. 

In  Science,1  however,  the  following  notice  concerning  Curtice's  obser- 
vation is  found: 

EARLY   STAGES    IN   THE   LIFE    OF   T^ENIA   PECTINATA. 

Thousands  of  sheep  and  lambs  perish  every  winter  on  the  ranches  west  of  the 
Missouri  Kiver.  They  are  not  apparently  afflicted  with  any  disease.  They  are  weak 
and  lean  in  the  fall,  and  simply  seem  to  be  unable  to  withstand  the  severity  of  the 
blizzards.  The  Bureau  of  Animal  Industry  of  the  Agricultural  Department  has  been 
engaged  in  an  investigation  to  ascertain,  if  possible,  the  cause  of  the  weakness  of 
the  animals  that  perish,  and  Mr.  Cooper  Curtice  visited  the  West  in  the  prosecution 
of  this  work.  An  examination  of  the  viscera  of  slaughtered  sheep  and  lambs,  fat 

1  March  23,  1888. 


200  TAPEWOEMS  OF  HARES  AND  RABBITS— STILES. 


and  healthy  ones,  as  well  as  those  that  were  weak  and  lean,  disclosed  the  fact  that 
they  were  almost  without  exception  infected  with  tapeworms,  which  were  found  in 
the  duodenum  and  gall  duct.  In  the  latter  they  were  frequently  so  numerous  as  to 
close  it  up,  and  cause  a  suspension  of  its  functions. 

For  the  purpose  of  continuing  his  studies  Mr.  Curtice  brought  from  the  West  a 
number  of  lambs,  which  were  killed  at  intervals  and  their  viscera  examined,  and 
this  material  having  been  exhausted,  and  it  being  inconvenient  and  expensive  to 
obtain  more,  he  turned  his  attention  during  the  past  winter  to  a  study  of  the  early 
stages  in  the  life  of  the  Tcenia  pectinata  (common  unarmed  tapeworms  of  the  rabbit). 
In  studying  these  Mr.  Curtice  thinks  he  has  made  some  interesting  discoveries, 
which  he  presented  to  the  Biological  Society  of  Washington  at  a  recent  meeting. 

The  variety  examined  is  found  abundantly  in  nearly  all  rabbits  in  this  locality. 
The  life-history  of  the  armed  tapeworms  of  man  and  dogs  has  long  been  written, 
but  that  of  the  unarmed  species  inhabiting  our  domestic  animals,  especially  cattle 
and  sheep,  is  as  yet  comparatively  unknown.  As  far  as  has  been  ascertained,  the 
life-history  of  the  Tcenia  pectinata  is  embraced  in  two  stages.  The  first  covers  the 
development  of  the  ova  into  the  embryo,  which  is  ready  to  leave  the  parent  Tcenia; 
the  other  covers  the  period  of  growth  from  the  youngest  forms  yet  found  in  rabbits 
to  the  adult  stage.  The  life  of  the  Tcenia  from  the  time  they  leave  the  first  rabbit 
as  an  embryo  until  they  are  found  as  young  Tcenia  in  the  second  rabbit  infected  has 
as  yet  been  unascertained.  Among  the  theories  that  have  been  advanced  is  one 
that  they  pass  this  stage  upon  the  ground,  are  eaten  by  insects,  snails,  or  crusta- 
ceans, and  that  these  are  then  eaten  by  the  rabbits.  This,  however,  is  only  a  theory, 
as  none  have  ever  been  found  in  snails,  insects,  or  crustaceans. 

It  was  Mr.  Curtice's  good  fortune  to  find  a  rabbit  which  had  recently  been  infected 
with  these  peculiar  parasites,  none  of  which  ^ere  over  3  centimeters  in  length, 
many  of  them  being  less  than  5  millimeters  long.  There  were  more  Twnia  in  that 
rabbit  than  any  he  had  ever  seen  before — about  85.  Among  the  smaller  Tania  were 
several  specimens  that  showed  the  stages  of  development  from  nonsegmented,  armed 
forms,  to  segmented,  unarmed  forms.  Mr.  Curtice  showed  "to  the  society  specimens 
illustrating  the  different  stages. 

The  youngest  forms  detected  were  not  the  smallest,  but  measured  about  one-half  a 
centimeter  in  length.  They  contained,  in  addition  to  the  four  suckers,  a  cup-shaped 
cavity  in  the  place  of  the  rostellum.  Around  the  border  of  this  cup-shaped  cavity 
were  situated  85  or  90  hooks.  The  older  specimens  show  a  similar  cavity  with  no 
hooks.  Still  older  ones  show  no  cavity  at  all.  All  of  these  were  in  the  nonseg- 
mented stages,  but  other  forms,  some  of  them  smaller,  were  without  signs  of  hooks, 
and  had  already  begun  segmentation. 

Mr.  Curtice  compared  these  stages  with  similar  stages  in  Twnia  seraia  [serrata~\, 
and  said  that  the  youngest  stage  of  the  Tcenia  pectinata  was  probably  a  cysticercoid 
stage  and  not  the  cysticercal,  and  that  this  was  indicated  by  the  cup-shaped  cavity 
in  the  youngest  forms  of  the  Tcenia  pectinata. 

In  discussing  the  classification  founded  on  the  presence  or  absence  of  hooks,  he 
declared  it  to  be  incorrect,  since  the  discovery  described  above  shows  that  the 
unarmed  species  in  adult  stages  are  armed  in  earlier  stages. 

The  speaker  exhibited  some  elegant  drawings  made  by  Dr.  George  Marx,  illus- 
trating the  embryo  as  it  leaves  the  parent  Tcenia.  This  embryo  is  six-hooked  and 
surrounded  by  a  curious  pyriform  envelope,  to  which  there  is  a  double  prolongation, 
surmounted  by  a  cap  of  the  same  substance.  This  cap  has  a  shredded  border,  and 
is  believed  to  be  the  remnants  of  a  mass  which,  in  an  earlier  stage,  completely  sur- 
rounded the  embryo.  This  peculiar  envelope  has  been  previously  noticed  in  Italy 
by  Perroucito  [Perroucito]  and  in  France  by  Raillet  [RaillietJ.  This  stage  is 
similar  to  that  found  in  Tcenia  expansa,  the  unarmed  tapeworm  in  sheep. 

This  article  was  copied  in  the  Texas  Live  Stock  Journal,1  but,  owing 

'April  14,  1888. 


NO.  1105.  PROCEEDINGS  OF  TIJE  NATIONAL  MUSEUM.  201 

to  the  inaccessibility  of  the  publications  and  the  lack  of  details,  Cur- 
tice's observation  has  not  received  much  attention  from  helmiu- 
thologists. 

The  only  authors — so  far  as  I  can  find — who  have  taken  cognizance 
of  it  are  Neumann,1  in  1892,  and  Kailliet,2  in  1893,  and  Braun.  Railliet 
writes  as  follows : 

Subfamily  Anoploceplialinw.  *  *  *  The  life  history  is  still  unknown.  However 
C.  Curtice  lias  made  an  interesting  observation  on  Lepua  sylvaticu*  t  which  will  possi- 
bly place  experimenters  in  a  position  to  determine  the  development.  He  found  in 
the  intestine  a  large  number  of  small  Twniadw  which  were  still  very  young,  but  in 
different  stages  of  development.  Some  of  them  5  mm.  long,  nonsegmented,  pos- 
sessed between  the  suckers  a  dome-shaped  depression,  bordered  with  85  to  90  hooks; 
others,  still  older,  had  lost  their  hooks,  while  some  did  not  even  show  the  cor- 
responding depression ;  finally,  some  of  them  were  segmented,  but  all  of  these  were 
unarmed.  One  is  thus  led  to  suppose  that  the  larval  stage  of  the  Anoplocephalinw  is 
represented  by  an  armed  cysticercoid  and  that  the  hooks  disappear  during  the 
development.  (Free  translation.) 

Curtice's  observation  I  confirmed  and  extended  in  1894.3  Since  pub- 
lishing this  note,  Hassall  has  found  the  same  young  stages  in  several 
rabbits  (L.  sylvaticus]  in  Maryland,  and  with  this  material  the  former 
description  can  be  amplified.  Of  the  young  forms  collected  some  were 
studied  fresh,  others  mounted. 

Unarmed  forms. — Nine  of  the  mounted  specimens  showed  no  trace 
of  any  rostellum  or  hooks,  but  on  the  other  hand  some  of  them  exhib- 
ited traces  of  segmentation.  The  details  of  measurements,  etc..  are  as 
follows : 

1.  0.544  mm.  long;   head,  0.24  mm.  broad  by  0.208  mm.  long;  constriction  back  of 
suckers  0.128  mm.  broad;  suckers,  0.112  mm.  in  diameter. 

2.  0.848  mm.  long;  head,  0.256  mm.  broad  by  0. 192  mm.  long ;  constriction  back  of 
head,  0.12  rum.  broad. 

3.  0.304  mm.  long;  head,  0.208  mm.  by  0.208  mm. 

4.  7  nun.  long;  head,  0.448  mm.  broad  by  0.32  mm.  long;  suckers,  0.196  mm.  diame- 
ter; constriction  back  of  head,  0.368  mm.  broad.     The  transverse  lines  of  the  seg- 
mentation become  indistinctly  visible  almost  immediately  back  of  the  head,  but  no 
genital  anlageu  are  seen  in  any  portion  of  the  specimen. 

5.  0.816  mm.  long;  head,  0.256  mm.  broad  by  0.24  mm.  long;  signs  of  segmentation. 

6.  0.64  mm.  long;  head,  0.224  mm.  broad  by  0.176  mm.  long;  suckers,  0.112  mm.  in 
diameter;  segmentation  begins  0.224  mm.  back  of  the  head. 

7.  0.816  mm.  long;  head,  0.24  mm.  broad  by  0.192  mm.  long;  segmentation  percep- 
tible, 0.4  mm.  back  of  head. 

8.  0.64  mm.  long;  head,  0.24mm.  broad  by  0.176mm.  long;  segmentation, 0.224 
mm.  back  of  head;  suckers,  0.144  mm.  in  diameter. 

9.  0.656  mm.  long;  head,  0.24  mm.  broad  by  0.16  rnm.long;  suckers,  0.112  mm.  in 
diameter;  segmentation  begins  0.288  mm.  back  of  head. 

Armed  forms. — Twenty-seven  mounted  specimens  in  which  rostellum 
and  hooks  were  present  varied  in  measurements  as  follows:  Length, 


1  Traite"  des  maladies  parasitaires,  2d  ed.,  p.  461. 

2  Trait6  de  Zool.  med.  et  agric.,  I,  p.  268. 

3  Notes  sur  les  Parasites— 31 :  Une  phase  precoce  du  tenias  du  Lapin,  Bull.  Soc. 
zool.  France,  XIX,  pp.  163-165. 


202  TAPEWOEMS  OF  HARES  AND  BABBITS— STILES.  VOL.XIX. 

0.560  to  1.800  mm. ;  head,  0.352  to  0.512  mm.  long  by  0.320  to  0.480  mm. 
broad;  rostellum,  0.176  to  0.240  mm.  long  by  0.112  to  0.160  mm.  broad; 
number  of  hooks  on  rostellum,  100  to  122;  on  some  fresh,  specimens  as 
few  as  90  hooks  were  counted;  size  of  hooks  on  rostellum,  18  to  24  // 
long. 

As  in  the  case  of  the  unarmed  heads  mentioned  above,  there  was  a 
general  though  not  absolute  agreement  between  the  size  of  the  scolex 
and  the  length  of  the  parasite,  and  there  is  no  question  in  my  mind 
that  the  head  of  a  tapeworm  is  subject  to  increase  in  size  after  entering 
its  final  host;  numerous  observations  upon  young  specimens  of  tape- 
worms from  sheep  support  this  view. 

In  none  of  these  armed  specimens  was  there  the  slightest  trace  of 
segmentation.  In  many  cases  the  armature  was  not  complete  either 
upon  the  suckers1  or  upon  the  rostellum,  but  in  all  cases  some  hooks 
were  found,  and  the  rostellam  was  always  visible. 

Taking  all  these  observations  into  consideration.  I  am  forced  to  the 
conclusion  that  the  unarmed  forms  and  the  armed  forms  represent  the 
young  stages  of  two  different  species.  The  unarmed  forms  I  am  in- 
clined to  bring  into  connection  with  C.  variabilis,  p.  192,  while  the  close 
agreement  between  the  rostellum  of  this  young  stage  with  that  of  D. 
salmoni,  the  agreement  in  the  size  of  its  hooks,  the  agreement  in  the 
general  arrangement  of  the  hooks  on  the  suckers,  the  fact  of  their 
presence  in  the  same  host  species,  and  finally  the  fact  that  one  of  the 
adult  specimens  ofD.salmoni  (No.  1124,U.S.N.M.)  was  found  in  the  same 
locality  in  which  these  forms  were  found,  all  lead  me  to  the  conclusion 
that  the  young  armed  stage  here  described  represents  the  young  of 
Davainea  salmoni.  Experimental  demonstration  of  this  view  is,  how- 
ever, lacking.  In  several  specimens  studied  alive,  the  cysticercoids 
were  surrounded  by  a  membrane  (Plate  XXV,  figs.  4,8,10),  which, 
however,  became  entirely  lost  upon  being  subjected  to  technique. 

Armed  larval  forms  distributed  as  follows. — America:  Collections 
Bureau  of  Animal  Industry;  U.S.N.M. ;  Harvard;  Leidy;  Hassall; 
Stiles.  Europe:  British  Museum  ;  E.  Blanchard ;  liailliet ;  Moniez; 
Zschokke;  Parona;  Berlin  Museum;  Leuckart;  von  Linstow:  Looss; 
Halle  Zoological  Institute;  F.  E.  Schulze.  Specimens  will  also  be  sent 
to  Neumann,  Stossich,  Monticelli,  and  Max  Braun. 

Diagnosis. — Davainea  salmoni,  Stiles,  1895.  Strobila  attains  86  mm. 
or  more  in  length  and  3  mm.  in  breadth,  and  contains  about  450  seg- 
ments which  vary  in  form  from  nearly  rectangular  to  infundibuliform 
according  to  contraction,  most  segments  being  much  broader  than  long, 
the  distal  15  to  20  segments  becoming  longer  and  narrower,  nearly 
square,  1.8  by  1.4  mm.  Head  0.6  to  0.736  mm.  broad  by  0.38  to  0.448 
mm.  long.  Rostellum  retractile,  0.1  to  0.14  mm.  in  diameter,  armed 

'The  number  of  hooks  given  for  the  suckers  (150  to  200  at  least)  in  Notes  sur  les 
Parasites — 31 — is  very  greatly  underestimated.  It  is  utterly  impossible  to  count  them, 
but  I  should  now  estimate  the  number  about  750  for  each  sucker. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  203 


with  a  double  row  of  hooks  20  /*  long,  about  GO  hooks  in  each  row. 
Suckers  comparatively  large,  0.25  mm.  in  diameter,  armed  with  numer- 
ous hooks  of  various  sizes,  the  prongs  of  the  longest  attaining  10  /4  in 
length.  These  hooks  are  arranged  in  about  7  to  9  concentric  rows  con- 
taining about  750  (estimated)  hooks  of  various  sizes.  Neck  short  and 
thin,  0.8  mm.  long  by  0.48  mm.  broad.  Anlage  of  the  genital  organs 
appears  about  the  one  hundred  and  seventieth  segment.  Genital  pores 
irregularly  alternate  situated  in  the  middle  to  near  the  proximal  por- 
tion of  the  margin,  cirrus  anterior  to  vagina;  cirrus  pouch  small,  0.12 
to  0.14  mm.  long  by  44  to  64  /<  broad,  muscular  layer  may  attain  12  to 
20  fj.  in  thickness.  Testicles  occupy  almost  the  entire  median  field 
except  the  center  portion  in  which  the  female  glands  are  located.  Eggs 
are  arranged  in  egg  capsules,  about  160  visible  from  one  surface  of  the 
segment  and  measuring  0.112  to  0.118  mm.  in  diameter;  3  to  15  ova 
measuring  20  to  24  /*  in  diameter  in  each  capsule. 

Hosts. — Eastern  Jackass  Hare  (Lepus  melanotis)  by  Tallichet  and 
Norgaard;  cottontail  (L.  sylvaticus)  by  Curtice,  Hassall,  and  Stiles; 
intermediate  host  unknown*.  Several  specimens  found  in  collection  of 
Leidy,  but  collector's  name  is  not  stated  on  label. 

Types. — Description  taken  from  four  strobile  with  heads,  and  several 
fragments.  Bureau  of  Animal  Industry  Cestode  series  No.  1196  desig- 
nated as  type  and  deposited  in  the  United  States  National  Museum. 
Paratypes  distributed  as  follows :  Collection  of  Bureau  of  Animal  Indus- 
try; Berlin  Museum;  collection  of  K.  Blanchard.  Fragments  to  South 
Kensington  Museum  and  Vienna  Museum.  Technique  of  type  specimen 
(poorly  preserved),  alcohol,  hiematoxylin. 

Geographical  distribution.— United  States  of  North  America;  Texas 
(by  Tallichet  and  Norgaard);?  (by  Curtice);  Maryland  (by  Hassall  and 
Stiles). 

GENERAL  REMARKS. 

/  Several  interesting  problems  arise  in  connection  with  the  adult  ces- 
todes  described  above  and  allied  forms,  to  which  it  may  be  well  to 
draw  attention,  notwithstanding  the  fact  that  the  data  at  hand  are 
not  so  complete  and  satisfactory  as  might  be  desired. 

The  uterus:  Comparing  the  uteri  of  the  known  forms  of  the  Anoplo- 
cephalinge  (exclusive  of  Stilesia),  we  find  at  least  three  distinct  types 
represented  in  both  the  single-pored  and  the  double-pored  genera. 

I.  The  most  simple  uterus  appears  first  as  a  simple  transverse  tube 
extending  across  the  median  field- of  the  segment,  as  in  Anoplocepliala. 
This  same  type  of  uterus  is  evidently  found  (see  Meyner)  also  in  Bertia 
mucronata  and  Bertia  conferta.  In  the  double-pored  genus  Cittotcenia 
the  anlage  is  generally  single, but  in  some  cases  (C.  variaMlis}  it  may 
be  single  or  double  (divided)  in  the  same  strobila. 

The  increase  in  the  size  of  this  type  of  uterus  takes  place  by  an 
expansion  of  the  uterus  at  certain  points,  giving  rise  to  proximal  and 
distal  blind  sacs  or  pouches.  These  pouches  may  be  very  distinct  and 


204  TAPEWORMS  OF  HAKES  AND  RABBITS—  STILES. 


well  defined,  as  in  Anoplocephala  mamillana,  A.  wimerosa,  Cittotccnia 
marmotce,  C.  pectinata  ;  or  they  may  be  less  distinct  (C.  variabilis,  C. 
prcecoquis)-,  or  they  may  be  totally  absent  (C.  variabilis),  in  which  case 
the  uterine  tube  appears  as  an  enlarged  sac.  In  some  species  where 
they  are  at  first  well  developed  (Anoplocephala  mamillana  and  A. 
wimerosa)  their  boundaries  may  disappear  in  the  older  segments,  the 
uterus  appearing  as  a  sac. 

The  uterus  of  C.  denticulata  seems  to  be  more  complex  and  requires 
further  study. 

II.  The  genus  Thysanosoma  presents  a  totally  different  uterus.    The 
anlage  is  originally  a  simple  transverse  tube  as  in  the  cases  just  men- 
tioned, but  the  increase  in  size  takes  place  by  two  different  methods, 
which  gives  rise  to  quite  a  complicated  structure.     First,  the  anlage 
increases  greatly  in  length,  giving  rise  to  numerous  folds  running  longi- 
tudinally with  the  worm;  next,  this  undulate  tube  forms  blind  pouches 
which  are  surrounded  by  a  heavy  layer  of  connective  tissue,  giving  the 
pouch  a  characteristic  appearance;   the  ova  are  deposited  in  these 
pouches  and  the  original  tube  remains  empty.    The  uterus  of  Stilesia 
globipunctata  bears  a  certain  resemblance  to  the  uterus  of  Thysanosoma 
in  that  the  egg  pouches  have  a  somewhat  similar  structure. 

III.  A  third  type  of  uterus  appears  to  be  presented  by  the  genera. 
Andrya  and  Moniezia,  and  this  type  will  require  considerable  study 
before  it  can  be  correctly  interpreted.     From  the  present  status  of  our 
knowledge,  however,   it  may  be  described  as  a  complex   system  of 
branching  and  anastomosing  tubes,  a  single  set  being  present  in 
Andrya,  a  double  set  in  Moniezia.     In  the  later  stages  of  the  uterus 
the  boundaries  of  the  tubes  disappear  and  the  uterus  appears  as  a  sac. 

A  transverse  uterine  anlage  has  not  yet  been  recorded  for  these 
genera,  except  in  the  case  of  Moniezia  trigonophora^  for  which  species 
Stiles  and  Hassall1  in  1893  have  figured  a  short  transverse  canal  which 
represented  the  earliest  uterine  stage  they  observed. 

An  interesting  problem  now  presents  itself,  namely:  In  what  relation 
do  these  types  stand  to  each  other?  Have  we  here  three  parallel 
series  of  originally  single-pored  cestodes,  each  series  with  a  particular 
type  of  uterus,  and  have  the  double  pores  been  acquired  independently 
after  the  three  types  of  uteri  became  established?  Or,  do  the  single- 
pored  forms  represent  one  branch  and  the  double-pored  forms  represent 
another,  and  have  the  three  different  types  of  uteri  been  developed 
independently  in  each  branch? 

The  former  supposition  appears  to  me  much  the  more  probable,  for 
in  Thysanosoma  giardi  and  Cittotcenia  variabilis  we  find  both  single  and 
double  pored  segments. 

This  case  represents  one  of  the  most  interesting  cases  of  homo- 
plasy  which  has  ever  been  observed  in  helminthology  and  should  be 


1  Revision  of  adult  cestodes  of  cattle,  sheep,  and  allied  animals,  Bull.  4,  Bur.  An. 
Ind.,  U.  S.  Dept.  Agric.,  pis.  vm,  fig.  5 ;  ix,  fig.  3. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  205 


studied  further  when  proper  material  is  found.  It  is,  however,  by  no 
means  the  only  case  of  homoplasy  brought  forward  in  helminthology. 
I  have1  elsewhere  called  attention  to  the  branching  of  the  intestine 
in  two  distinct  types  of  flukes,  as  represented  by  Fasciola  liepatica  and 
Distomum  delpliini.  The  presence  of  more  than  two  testicles,  as  in 
two  or  more  other  distinct  types  of  flukes  represented  by  Pleorchis 
polyorcUis  and  P.  mollis  on  the  one  hand  and  Distomum  cycjnoides  or  Z>. 
richarcU  on  the  other  is,  in  iny  opinion,  another  case  of  homoplasy. 
The  development  of  double-pored  forms  in  two  distinct  families  (the 
Bothriocephalkhe — Blanchard's  genus  Krabbea,  and  a  number  of  cases 
in  which  a  few  double-pored  segments  have  been  found  in  Bothrio- 
cephalm  latm — and  the  Taeniidue  forms  a  fourth  case  of  the  same  nature. 

One  of  the  greatest  criticisms  upon  the  present  classification  of  para- 
sites seems  to  rne  the  fact  that  systematists  have  not  taken  into  con- 
sideration the  principle  so  well  established  in  paleontology  and  other 
branches  of  zoology,  namely,  that  the  same  structure  may  develop 
independently  in  several  parallel  series  of  animals.  The  sooner  this  prin- 
ciple is  acknowledged  the  sooner  we  shall  have  a  natural  classification. 

Zone  of  lateral  growth. — A  comparison  of  the  older  and  younger  seg- 
ments of  numerous  Ta3uiida3  shows  that  as  the  segments  grow  broader, 
the  median  field — namely,  the  space  between  the  two  ventral  canals — 
increases  much  more  rapidly  in  proportion  than  the  lateral  fields — 
namely,  the  space  between  the  ventral  canals  and  the  lateral  mar- 
gin. This  establishes  the  median  field  as  the  greatest  zone  of  lateral 
growth.  No  particular  narrow  zone  of  the  median  field  can  be  looked 
upon  as  the  zone  of  lateral  growth  for  all  cestodes,  as  is  shown  by  a 
comparison  of  a  large  number  of  forms.  In  Tccnia  (type  T.  solium], 
for  instance,  the  entire  median  field  appears  to  participate  in  a  more  or 
less  uniform  lateral  growth,  for  the  genital  glands  retain  their  rela- 
tive position,  increasing  in  size  in  proportion  to  the  increase  in  size 
of  the  segment;  the  same  appears  to  hold  for  Davainea.  In  the 
anoplocephaline  forms  we  find  some  interesting  variations  in  the  zone 
of  growth,  as  is  shown  by  the  relative  position  of  the  ovaries  to  the 
ventral  canals  or  to  the  median  line.  In  Thysanosoma  yiardi  the  ovary 
bears  an  almost  constant  relation  to  the  ventral  canal,  while  the  dis- 
tance between  the  ovary  and  the  median  line  constantly  increases  as  the 
segment  grows  broader;  in  this  case,  therefore,  there  is  but  little  growth 
between  the  ovary  and  the  canal,  while  the  growth  in  the  median  side 
of  the  ovary  is  very  marked.  The  same  holds  true  to  a  lesser  degree 
in  Bertia  americana.  In  Moniezia  expansa  and  M.  pianissimo,  also  the 
relation  of  the  ovaries  to  the  ventral  canals  remains  almost  constant, 
while  the  chief  zone  of  growth  is  in  the  median  field  between  the 
ovaries. 

'The  anatomy  of  the  large  American  fluke  (Fasciola  magna]  and  a  comparison 
with  other  species  of  the  genus  Fasciola  s.  st.  vide  p.  221,  Jour.  Comp.  Med,  Vet. 
Arch.,  1895. 


206  TAPEWORMS  OF  HARES  AND  RABBITS— STILES. 


Iii  Cittotcenla  marmotce  (Plate  XI,  fig.  5),  C.  prcecoquis,  and  C.  pecti- 
nata  (Plate  XVI),  on  the  other  Land,  there  also  is  a  marked  lateral 
growth  between  the  ovary  and  the  ventral  canal  while  the  lateral  field 
remains  nearly  the  same  width  throughout  the  entire  length  of  the 
strobila.  In  C.  variabilis  (Plates  XIX-XX)  there  is  a  marked  growth 
in  the  lateral  fields. 

Abnormalities  in  the  segments. — Eiehm  has  already  recorded  segments 
of  Ciitotcenia  denticulata  with  three  sets  of  female  glands,  and  I  have 
recorded  segments  of  C.  variabilis  angusta  with  a  single  pore.  New 
cases  of  both  of  these  variations  have  been  noticed  in  the  preparation 
of  this  paper,  namely,  triple  sets  of  female  glands  in  C.  denticulate 
and  single  pores  in  C.  variabilis.  I  can  look  upon  these  cases,  however, 
only  as  variations,  similar  to  the  occasional  double  pores  in  Bothrioceph- 
alus  latus  and  Thysanosoma  (jiardi,  and  can  not  ascribe  to  them  any 
particular  value  from  a  systematic  standpoint. 

The  value  of  the  genital  pores  in  classification. — Objection  has  arisen 
in  some  quarters  to  adopting  the  genital  pore  as  the  basis  of  classifica- 
tion. I  both  agree  and  disagree  with  this  objection.  It  must  not  be 
forgotten  that  when  Blanc-hard  used  the  genital  pores  as  basis  for  his 
classification,  he  naturally  used  the  pore  as  representative  of  the  geni- 
tal system;  thus,  if  the  pores  are  single,  the  entire  female  system  is 
generally  single;  if  the  pores  are  double,  the  entire  system  of  female 
glands  is  generally  double.  Viewed  from  this  standpoint  (which  is  the 
only  logical  interpretation  I  can  give  to  Blanchard's  classification,  pro- 
posed in  1891),  I  must  insist  upon  the  great  value  of  the  pores  in  classi- 
fying cestodes.  At  the  same  time  I  fully  agree — and  have  stated  so 
more  than  once  before — that  the  pores  un associated  with  other  charac- 
ters can  not  be  relied  upon  as  basis  for  a  natural  classification  of 
cestodes.  Internal  topographical  anatomy  must,  in  my  opinion,  form 
the  basis  of  the  natural  classification  for  both  Oestoda  and  Trematoda. 
The  size  and  form  of  the  segments  are  characters  which  we  should  use 
with  the  greatest  caution,  always  making  due  allowances  for  technique 
and  contraction ;  as  generic  characters  I  can  under  no  circumstances 
admit  their  validity,  and  must  therefore  reject  Sonsino's  recently  pro- 
posed genus  Panceria  (based  essentially  upon  double  pored  segments 
which  are  longer  than  broad)  unless  other  characters  are  brought  for  ward 
to  place  the  genus  on  a  firmer  footing. 

In  systematic  work  in  helminthology  we  must  not  lose  sight  of  another 
principle  which  is  well  acknowledged  in  other  specialities,  namely,  that 
a  given  character  which  may  be  of  great  importance  in  classifying  the 
species  or  genera  of  one  group  does  not  necessarily  hold  as  a  taxo- 
nomic  character  in  all  genera  or  higher  groups  of  the  same  order  or 
class.  Thus  the  unilateral!  ty  of  the  pores  in  Hymenolepis  and  Anoplo- 
cepliala  appears,  so  far  as  investigations  have  gone,  to  be  a  very  impor- 
tant and  constant  character,  while  the  same  character  must  be  used 
with  the  utmost  precaution  in  the  genus  Davainea.  I  have  shown 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  207 


above  that  the  pores  of  I),  salmoni,  for  instance,  may  be  either  unilat- 
eral or  irregularly  alternate;  the  same  holds  for  J).  tetragona,  and 
apparently  also  for  Andrya.  The  characters  to  be  used  in  classify- 
ing the  species  of  any  given  genus  must  be  determined  separately  for 
every  genus  by  an  examination  of  a  large  series  of  specimens  from  dif- 
ferent localities  and  from  different  hosts.  The  unsatisfactory  condition 
in  which  I  have  been  obliged  to  leave  the  genera  Bertia  and  Andrya 
is  directly  attributable  to  the  limited  amount  of  material  at  my  disposal, 
and  the  fact  that  what  few,  and  for  the  most  poorly  preserved,  specimens 
I  have  examined  represent  only  a  few  localities  and  a  few  hosts. 

The  influence  of  a  host  upon  its  parasites. — Considerable  has  been 
written  upon  the  influence  of  various  parasites  upon  their  frosts,  but 
very  few  authors  have  ventured  to  publish  upon  the  influence  of  the 
hosts  upon  the  parasites.  This  is  probably  due  to  two  reas.ons:  first, 
to  the  tendency  on  the  part  of  many  helmiuthologists  to  take  the  host 
species  as  a  specific  character  of  the  parasite,  and  secondly  the  fact 
that  the  influence  exerted  by  the  parasite  upon  its  host  is  a  matter  of 
economic  importance,  while  the  influence  of  the  host  upon  the  parasite 
would  be  looked  upon  by  most  workers  as  purely  of  scientific  interest. 
To  deny  the  economic  importance  of  the  host  influence  upon  the  parasite 
is,  however,  to  my  mind  a  short-sighted  policy,  for  it  must  be  this  very 
host  influence  (i.  e.,  environment)  which  has  played  an  important  role  in 
the  evolution  of  species,  and  which  must  have  resulted  in  differentiating 
species  and  varieties,  each  of  which  after  a  time  becomes  in  some  cases 
at  least  dependent  upon  a  single  host.  The  genera  Demodex,  Psoroptes, 
and  Cfiorioptes  form  excellent  examples  of  the  point  under  considera- 
tion; in  these  genera  we  find  varieties  which  resemble  each  other  so 
closely  that  it  is  often  difficult  and  even  impossible  to  determine  the 
variety  without  knowing  the  host;  experiments  to  breed  some  of  the 
varieties  of  these  genera  upon  other  than  their  regular  hosts,  even 
though  the  experiment  animal  harbors  a  very  closely  allied  variety  of 
the  same  species,  have  been  totally  negative.  This  same  variation  is 
noticeable  among  worms.  Specimens  of  Moniezla  expansa  taken  from 
Ovis  aries  are  for  instance  totally  different  from  the  specimens  of 
the  same  species  found  in  Ovis  laticauda;  the  strobilse  from  this  latter 
host  recently  studied  by  Setti,  which  I  have  been  able  to  examine 
through  the  kindness  of  my  Italian  colleague  Prof.  0.  Parona,  can  not 
I  believe  be  separated  specifically  from  the  European  form,  yet  they 
certainly  represent  a  distinct  variety.  Lonnberg  has  called  attention 
to  the  variation  in  the  hooks  of  avian  cestodes  taken  from  different 
hosts.  I  have  also  repeatedly  noticed  a  variation  in  the  spicules  of  the 
same  species  of  nematode  (Strongylus  contortus,  for  instance)  taken  from 
cattle  and  sheep.  To  clearly  define  the  varieties  peculiar  to  certain 
hosts  touches  directly  upon  the  question  of  transmission  and  infection, 
and  on  this  account  I  submit  that  the  study  is  of  great  economic  as 
well  as  scientific  importance. 


208  TAPEWOEMS  OF  HAKES  AND  RABBITS— STILES. 


The  division  of  the  genus  Tcenia. — In  connection  with  the  citations, 
p.  162,  from  Meyner's  work,  I  take  this  occasion  to  reply  to  his  remarks 
regarding  the  efforts  which  E.  Blancliarcl,  Eailliet,  Hassall,  and  I  have 
recently  made  to  bring  some  order  into  the  genus  Tcenia.  Upon  page 
8  of  his  thesis  he  says : 

Die  Versucbe  von  Blauchard,  Stiles  und  Railliet  die  Uebersichtlichkeit  de'r 
Familio  der  Tamiaden  durcli  Zerlegung  in  eine  Auzahl  von  Unterfainilien  zu  erleieb- 
tern,  siud  uiclit  als  besonders  gliicklicbe  zu  bezeicbiien,  namentlicb  mit  Riicksicbt 
darauf,  dass  die,  von  verscbiedeuen  Gesicbtspunkten  ausgehende,  stets  nur  an  einer 
bescbrankten  Anzabl  von  Species  vorgenonimene  Eintbeilung  notbgedrungeu  aucb 
zu  verscbiedenen  Resultaten  fiibreu  musste.  Bevor  nicbt  durcb  eiugebende  For- 
schungen  wenigstens  die  typiscben  Hanptforinen  als  feststebend  angeseben  wer- 
deu  koimeri,  diirften  derartige  Experimeute  wobl  uicbt  auf  allgemeine  Anerkennung 
zu  recbnen  baben,  bis  dabiu  aber  verfeblen  sie  ganz  entscbiedeu  ibren  eigentlichen 
Zweck,  niimlicb  Klarbeit  nnd  Eiufacbbeit  in  die  Systematik  zu  briugen. 

In  writing  this  critique,  which  is  of  course  welcomed  both  by  my 
French  colleagues  Blanchard  and  Eailliet,  and  by  Hassall  and  myself  as 
showing  the  view  which  Dr.  Meyner  takes  of  our  work,  the  author  has 
unfortunately  overlooked  several  very  important  points.  First  of  all, 
he  has  overlooked  that  very  important  principle  so  well  expressed  by 
Bacon  in  the  words,  "Truth  emerges  sooner  from  error  than  from  con- 
fusion." iNone  of  us  look  upon  our  efforts  to  classify  these  parasites  as 
perfect,  free  from  criticism,  and  final  beyond  revision.  But  errors 
which  we  may  have  committed  in  these  attempts  to  aid  in  reducing 
the  chaos  which  exists  in  the  genus  Tcenia  can  be  corrected  very  easily. 
We  have  the  satisfaction  of  knowing  that  our  work  has  been  adopted 
by  a  number  of  specialists  in  helmiuthology  and  hardly  feel  called  upon 
to  apologize  for  the  attempts  we  have  made,  even  if  not  approved  of  by 
Veterinarian  Meyner. 

A  second  error  Dr.  Meyner  has  fallen  into  is  that  he  has  forgotten 
that  every  classification  must  undergo  an  evolution  (with  epigenesis !). 

He  would  evidently  have  us  include  all  four-suckered  tapeworms  in 
the  heterogeneous  and  collective  genus  Tcenia,  "until  at  least  the  typical 
chief-forms  can  be  looked  upon  as  established,"  although  he  failed  to 
give  an  explanation  of  what  he  meant  by  "the  typical  chief-forms."  If 
he  refers  to  the  type  species  of  genera,  he  has  lost  sight  of  the  fact  that 
the  type  species  of  a  genus  is  designated  by  any  given  author,  and  can 
not  be  changed  after  once  being  so  designated.  The  type  species  is, 
therefore,  optional,  subject  of  course  to  certain  principles,  in  the  case 
of  the  author  who  designates  it  and  obligatory  in  the  case  of  other 
authors.  Possibly  Meyner  refers  to  the  most  highly  specialized  species 
in  the  different  groups.  If  so,  however,  his  advice  to  wait  until  we 
study  up  such  forms  and  decide  which  are  the  most  highly  specialized 
is  wanting  in  reason.  It  is  certainly  far  better  to  create  a  new  genus 
for  a  form  when  we  can  not  logically  unite  it  with  the  known  genera, 
than  it  is  to  place  it  with  forms  with  which  it  does  not  agree  in  any 
important  character. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  209 

Of  course  our  efforts  at  classification  are  experimental — we  all  admit 
that;  but  from  the  very  nature  of  things  all  efforts  at  classification  in 
practically  unknown  groups  are  and  must  be  experimental.  The  classi- 
fications must  be  changed  time  and  again  as  new  facts  are  discovered. 
Nor  have  our  experiments  (or,  as  Meyner  puts  it,  "derartige  Experi- 
inente")  counted  upon  immediate  general  recognition  (allgemeine  Aner- 
kenuuug) ;  it  was  not  with  that  end  in  view  that  we  published  them. 
We  do  not  expect  to  see  our  proposed  classifications  adopted  by  zoolo- 
gists at  large  until  they  have  stood  the  test  of  other  specialists  in 
helminthology.  We  have  not  heard  as  yet,  however,  of  any  marked 
disapproval  of  the  genera  proposed  from  workers  who  were  acquainted 
with  the  forms  and  who  were  competent  to  pass  judgment  on  the  case. 
When  such  authors  propose  a  better  classification,  they  can  certainly 
count  on  IHauchard,  Eailliet,  Hassall,  and  myself  as  four  helmintholo- 
gists  who  are  ready  to  follow  them.  At  present,  however,  I  maintain 
that  the  classification  originally  proposed  by  Blanchard  and  since  that 
time  considerably  expanded  by  Eailliet,  Hassall,  and  myself  is  a  far 
more  natural  and  satisfactory  classification  oi  the  forms  treated  than 
any  other  classification  ever  proposed  for  the  same  forms.  I  am  fully 
convinced,  after  a  study  of  several  thousand  specimens,  that  the  main 
features  of  the  proposed  division  will  stand,  although  the  details  of  the 
system  may  undergo  some  changes.  Helminthologists,  as  a  class,  are 
ultra-conservative  in  every  line  except  species-making — and  yet  as  long 
as  the  Eudolphi-Diesing  school  exerts  such  a  powerful  influence  in  wield- 
ing the  yardstick  instead  of  the  microscope,  perhaps  this  generic  con- 
servatism should  be  looked  upon  as  a  blessing. 

A  third  error  of  Meyner's  is  that  he  does  not  understand  the  views 
which  he  has  attempted  to  criticise,  or  the  relative  rank  of  the  groups 
proposed,  and  he  ascribes  to  authors  propositions  which  they  never 
made.  Thus  he  states  (page  6) : 

Diese  Anoplocephalinen  theilt  er  (R.  Blanchard)  dann  init  Riicksicht  auf  die  Anord- 
nung  der  Geschlecktsorgane  in  3  Unterfamilien  [!]  ein  und  zwar  (1)  Genre  Moniezla 
*  *  * ;  (2)  Genre  Auoplocephala  *  *  * ;  (3)  Genre  Bertia 

Meyner  thus  makes  the  terms  subfamily  and  genus  synonymous — 
rather  a  novel  idea  in  systematic  zoology;  he  accredits  (page  8)  Blauch- 
ard  and  Eailliet  with  a  family  "Anoplocephalen,"  although  he  states 
a  few  lines  before  that  Eailliet  accepted  "Anoplocephaline"  as  a  sub- 
family. Upon  the  same  page  he  speaks  of  Bertia  as  a  genus  and  Cteno- 
tcenia  and  Andrya  as  "Arten."  It  does  not  seem  to  me  at  all  strange 
that  our  efforts  should  "fail  totally  in  their  object"  with  a  worker  who 
confounds  such  terms  as  species,  genus,  subfamily,  and  family. 

A  fourth  error  into  which  Meyner  has  fallen  in  the  passage  quoted 
is  the  assumption  that  we  have  taken  only  a  few  species  into  consid- 
eration in  making  our  classifications.  True,  we  have  not  felt  called 
upon  to  give  a  list  of  all  the  species  of  cestodes  with  which  we  have 
acquaintance,  either  through  personal  study  or  through  the  publications 
Proc.  N.  M.  vol.  xix 14 


210  TAPEWORMS  OF  HARES  AND  RABBITS—  STILE '£. 


of  our  colleagues,  but  it  would  be  going  a  little  far  to  assume  that 
we  did  not  have  other  forms  in  mind  at  the  time  we  revised  a  por- 
tion of  the  species  known  to  us.  Consistency  being  such  a  virtue, 
however,  if  Meyner  considers  the  species  mentioned  in  the  former 
papers  by  the  authors  he  criticises  as  representing  the  sum  total  of 
the  species  known  to  them,  he  will  probably  not  object  to  applying  the 
same  criterion  to  himself.  As  he  has  published  original  work  on  only 
two  species,  however,  it  might  be  difficult  for  him  to  justify  himself  in 
attempting  upon  the  basis  of  these  two  species  to  overthrow  the  con- 
clusions which  have  been  based  upon  a  study  of  a  much  larger  number 
of  forms  by  four  different  specialists,  all  of  whom  agree  in  the  general 
arrangement  and  differ  with  each  other  only  in  matters  of  detail. 
Meyner's  remarks  represent  an  excellent  example  of  what  is  so  com- 
mon in  science  to-day,  namely,  a  tendency  to  jump  at  conclusions  and 
to  generalize  in  a  too  dogmatic  manner  when  one  is  not  thoroughly  at 
home  on  the  subject  he  has  under  discussion. 

The  above  remarks  are,  I  think,  sufficient  to  show  that  Meyner's 
amusing  criticisms  would  better  have  been  submitted  to  more  careful 
thought  before  they  were  published.  They  can  not  have  much  weight 
with  helminthologists,  and  should  not  have  much  weight  with  zoolo- 
gists in  other  specialities. 

A  number  of  genera  not  discussed  in  this  paper  have  been  proposed 
by  authors  for  various  forms  in  the  family  Tasniidse.  Some  of  these 
genera  must  be  rejected.  In  regard  tofsome  of  the  others,  judgment 
must  be  reserved  for  the  present.  I  hope,  however,  to  publish  before 
long  a  summary  of  all  the  genera  proposed.  Regarding  the  newly  pro- 
posed genera  for  avian  cestodes,  see  Stiles,1  1896. 

Exchange  of  cotypes. — There  seems  to  be  a  popular  impression  among 
workers  that  it  is  a  perfectly  easy  matter  for  a  scientist  to  read  a 
description  of  a  species  or  genus  and  judge  of  the  validity  of  the  pro- 
posed form  without  examining  specimens.  This  impression  is  certainly 
true  in  some  cases,  but  must  always  be  taken  cum  grano  salis.  I  do  not 
hesitate  to  assert  that  not  one-half  of  the  species  and  genera  of  parasitic 
worms  ever  described  can  be  rightly  judged  from  their  descriptions,  nor 
is  this  always  attributable  either  to  the  description  or  to  the  ability  of 
the  worker  attempting  to  pass  judgment  upon  the  case.  It  is  rather 
attributable  to  the  undeniable  fact  that  a  person  obtains  an  entirely 
different  impression  from  a  study  of  the  objects  from  what  he  obtains 
from  reading  a  description,  be  it  ever  so  detailed  and  complete.  Many 
a  species  or  genus  has  been  accepted  or  rejected  by  an  author  who  would 
have  decided  differently  if  he  could  have  examined  cotypes  of  the 
forms  he  was  discussing. 

In  view  of  these  facts,  which  I  am  confident  the  specialists  in  hel- 
minthology  will  admit,  I  wish  to  appeal  again  to  helminthologists  to 


1  Report  upon  the  present  knowledge  of  the  tapeworms  of  poultry,  Bull.  12,  Bureau 
Animal  Industry,  Washington,  D.  C. 


NO.]  105.  PROCEEDINGS  OF  THE  RATIONAL  MUSEUM.  211 

deposit  cotypes  with  the  chief  museums  of  the  world.  It  has  been  our 
policy  to  do  this  as  far  as  possible,  and  as  soon  as  the  international 
postal  laws1  are  amended  so  as  to  allow  the  acceptance  of  these 
specimens  in  the  international  mails,  the  system  will  be  extended. 

As  a  matter  of  justice,  I  believe  the  type  specimen  belongs  in  the 
national  museum  of  the  country  in  which  the  specimen  was  collected. 
The  first  duplicate  (paratype)  belongs  by  right  to  the  Berlin  Museum,  as 
that  museum  now  possesses  more  types  of  parasitic  worms  than  prob- 
ably any  other  collection  in  the  world.  I  would  set  forth  the  claims 
of  the  United  States  National  Museum  for  the  second  duplicate  on  the 
grounds  that  the  European  workers  all  have  easy  access  to  the  speci- 
mens at  Berlin,  while  the  frequent  sending  of  a  rare  and  valuable 
specimen  from  Europe  to  this  continent  is  attended  with  more  or  less 
danger  of  loss.  Japan  certainly  has  a  claim  to  the  third  duplicate. 

It  is  indeed  time  that  helminthologists  give  the  question  of  the  pres- 
ervation of  types  their  serious  consideration.  Many  of  the  older  col- 
lections can  not  be  traced  5  Davaine's  collection  is  evidently  destroyed, 
and  many  more  private  collections  will  follow  it  if  more  attention  is  not 
given  to  this  subject. 

CONCLUSIONS. 

The  chief  results  of  this  paper  may  be  briefly  summarized  as  follows: 

1.  The  time  has  come  when  helminthologists  must  donate  their  type 
specimen  of  every  species  to  museums  where  they  can  be  properly  cared 
for  and  where  they  may  be  open  to  the  inspection  of  other  specialists. 
I  maintain  that  the  original  type  rightfully  belongs  to  the  national 
museum  of  the  country  in  which  the  specimen  was  collected ;  that  the 
second  specimen  (first  duplicate— first  paratype)  rightfully  belongs  to  the 
Berlin  Museum ;  that  the  United  States  National  Museum  has  a  valid 
claim  for  the  third  specimen  (second  duplicate),  and  that  Japan  has  a 
valid  claim  for  the  fourth  specimen. 

2.  In  order  to  insure  the  preservation  of  types  and  cotypes,  I  invite 
the  specialists  in  helminthology  to  unite  upon  some  regular  plan  for 
the  deposit  and  exchange  of  such  specimens. 

3.  Permanent  mount  in  balsam  appears  to  me  to  be  a  much  better 
method  of  preserving  a  type  than  to  retain  it  as  an  alcohol  specimen. 

4.  The  known    adult  leporine  cestodes  belong  to  the  five  genera: 
Anoplocephala,  Andrya,  Bertia,  Cittotcenia,  and  Davainea. 

5.  Of  these  genera  Anoplocepliala,  Cittotoenia,  and  Davainea  are  con- 
sidered as -perfectly  valid  genera;  Andrya  and  Bertia  are  left  subjiidice, 
although  it  is  my  impression  that  they  will  finally  be  recognized  as 
valid,  as  soon  as  sufficient  material  can  be  properly  studied. 

'Owing  to  the  absurd  international  postal  regulation  (16.  3.  b.)  excluding  zoological 
specimens  from  the  international  mails,  some  of  the  paratypes  mentioned  as  "  distrib- 
uted" in  this  paper  have  not  yet  been  sent  to  the  authors  named;  they  will  be 
forwarded  as  soon  as  an  opportunity  presents  itself. 


212  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 

6.  The  microscope  must  supplant  the  yardstick  and  internal  anatomy 
must  take  the  place  of  external  form  in  judging  the  validity  of  cestode 
and  trematode  genera  and  species. 

7.  The  principle  of  homoplasy  must  be  recognized  by  helmintholo. 
gists  as  well  as  by  other  zoologists,  and  any  classification  which  leaves 
this  important  and  well-recognized  principle  out  of  account  can  be  taken 
only  as  a  preliminary  (although  often  necessary)  study  (p.  204). 

8.  The  median  field  of  the  Tseniidie  is  the  seat  of  the  most  active  lat- 
eral growth,  and  the  same  rule  will  probably  be  found  to  apply  to  other 
families  of  Cestoda.    No  particular  longitudinal  zone  of  the  median  field 
can,  however,  be  named  as  the  zone  of  most  active  growth  in   all 
Ta3niida3  (p.  205). 

9.  The  armed  young  cestode  which  I  mentioned  in  Note  31 [  (1895)  is 
not  the  young  of  an  anoplocephaline  tapeworm,  as  Curtice,  Braun, 
Kailliet,  Neumann,  and  I  had  inferred,  but  represents  the  young  stage 
of  the  single-pored  cestode  referred  to  in  my  paper  in  1895.2    It  evi- 
dently belongs  to  Davainea  salmoni. 

10.  The  double-pored  cestode  with  occasional  single  pores,  described 
in  my  paper  in  1895,3  is  Cittotcenia  variabilis  angusta. 

11.  I  have  also  found  some  very  young  stages  of  an  unarmed  cestode 
in  the  intestine  of  the  cottontail  rabbit,  which  probably  belong   to 
Cittotcenia  variabilis.     This  young  stage  corresponds  to  what  we  may 
expect  to  find  as  the  larval  form  of  Moniezia  expansa  of  cattle  and  sheep, 
and  I  doubt  whether  it  will  be  possible  to  distinguish  it  from  the  young 
of  that  species.     This  renders  the  question  of  the  origin  of  the  tape, 
worms  of  cattle  and  sheep  more  complicated  than  it  was  formerly  sup- 
posed to  be,  and  demands  the  strictest  experimental  proof  on  the  part 
of  any  author  who  suspects  that  he  has  solved  the  mystery  of  the  life 
history  of  the  cestodes  of  cattle  and  sheep. 

12.  The  head  of   a    cestode   increases  in   size  after  the  parasite 
reaches  its  final  host,  as  is  shown  by  a  comparison  of  the  younger  speci- 
mens. 

13.  None  of  the  adult  leporine  tapeworms  thus  far  described  in  Europe 
have  as  yet  been  found  in  America.      The  American  forms  which  have 
been  published  as  "  Tccnia  pectinata"  must  be  distributed  over  several 
species  typical  to  this  continent. 

14.  The  following  table  includes  all  of  the  genera  at  present  recog- 
nized in  the  subfamilies  Ta3niin»,  Mesocestoidime,  Anoplocephalinse,  and 
Dipylidiinse  of  the  family  Ta3niidae.    A  number  of  other  genera  have 
been  proposed,  but  some  of  them  must  fall  as  synonyms,  while  judg- 
ment upon  others  must  be  reserved.     Several  of  the  genera  in  this  key 

1  Notes  sur  les  Parasites — 31:    Une  phase  prdcoce  du  Teuias  <lu  Lupin,  Bull.  Soc, 
zool.  France,  XIX,  1895,  pp.  163-165. 

2  Notes  on  Parasites — 36:    A  double-pored   cestode  with  occasional  single  pores, 
Centralbl.  f.  Bakteriol.  u.  Parasitenk.,  1.  Abt.,  XVII.  1895,  pp.  457-459. 

3Loc.  cit. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  213 


are  accepted  only  provisionally,1  while  Fimbriaria  is  used  at  present 
only  as  a  matter  of  convenience. 

Key  to  the  genera  of  the  subfamilies  TtBuiinaB,  MesocestoidinaB,  Ano- 
plocephalime,  and  Dipylidiinse  of  the  cestode  family  Tseniidie,  including 
all  the  species  of  adult  tapeworms  found  in  hares  and  rabbits. 

Tceniidce. 

[Tapeworms  of  liares  and  rabbits  in  roman.] 

1.  Head  with  four  cup-shaped  suckers;  rostellum  present,  but  not  always  evident; 

uterus  without  any  special  pore ;  genital  pores  generally  marginal 2 

Head  generally  absent;  genital  pores  marginal.     Host:  Birds 34 

2.  Genital  pores  veutro-median  (Mesocestoidina)) Mesocestoides.^ 

Genital  pores  marginal 3 

3.  Head  never  provided  with  hooks;  uterus  transverse  or  reticulate ;  egg  generally 

with  pyriform  body;  larval  stage  not  known;  adults  in  herbivorous  and  fru- 

givorous  mammals Anoploccplialina?  (p.  149),  5 

Head  nearly  always  provided  with  hooks  (T.  saginata  an  exception;   see  also 
Hymenolepis) 4 

4.  Egg  with  thin  outer  shell  and  thick  brown  inner  shell;  uterus  median  and  longitudinal 

with  lateral  branches;  larval  stage  a  cyslicercus,  cocnurus,  or  echinococcus,  generally 
in  herbivora;  adults  in  carnivora ( Tceniinw)  Tcenia.'-1 


'For  Andrya  and  Bertia,  see  pp.  154,  160  of  this  paper.     For  the  newly  proposed 
genera  Amabilia,  Choanotcenia,  Cotugnia.  JJicranotamia,  I)repanidota>,nia\v\  the  subfamily 
Dipylidiinse,  see  my  Report  upon  the  Present  Knowledge  of  the  Tapeworms  of  Poultry, 
Bull.  12,  Bureau  of  Animal  Industry,  1896. 
*  1863,  Hesocestoides,  VAILLANT,  L'Institut,  No.  1524.     Type,  Mesocestoides  ambiguus, 

Vaillant,  1863. 
1885,  Pti/chophysa,  HAMANN,    Zeitschr.  f.  wiss.  Zool.,  XLII,  p.  740.     Type,  Tcenia 

lineata,  Goeze,  1782. 
3  1758,  Twnia,  LINNAEUS,  Systema  naturae,  10  ed.,  p.  646,  819.     Type  by  elimination 

Tawia  solium,  Linmvus,  1758. 
1800,  Ali/selminthus,  ZEDEH,  Erster  Nachtrag  zur  Natnrg.  d.  Eingeweidewiirmer,  p. 

221.     Tamia,  Linnaeus,  renamed,  hence  type  Tania  solium,  Linmeus,  1758. 
1803,  Halt/sis,  ZEDER,  Anleitung  zur  Naturg.  d.  Eingeweidewiirmer,  p.  298.     Alysel- 

minthus,  Zeder,  renamed,  hence  type  Tcenia  solium,  Linmeus,  1758. 
1850,  Arhyncliotwnia,  DIESING,  Systema  helminthum,  I,  p.  497.     Subgenus  containing 

type  of  genus,  hence  type  Ta>nia  solium,  Linnrous,  1758. 
1858,  Acanthotrias,  WEINLAND,  Human  Cestoides,  p.  51.     Type,  Cysticercus  acantho- 

trias,  Weinland,  lSoS  =  Ciislicercuscellnlosw,  Rudolphi^:larva  of  Twnia  solium. 
1858,  Tamiarhyncltns,  WEINLAND,  Human  Cestoides,  p.  46.    Type,  Tcenia  mediocanellata, 

Kiichenmeister,  1852  =  Tcenia  saginata,  Goeze,  1782. 
1858,  Echinococcifer,  WEINLAND,  Human  Cestoides,  p.  52.     Type,  Tania  echinococcus, 

Siebold,  1853. 
1863,  Cystotwnia,  LEUCKART,  Die  menschlichen  Parasiten,  I,  p.  223.     As  subgenus 

including  type  of  genus,  hence  type  Tcenia  solium,  Linnaeus,  1758. 
To  this  list  of  synonyms  must  be  added  the  subgenera  Cysticercus,  Cocnurus,  and 
Echinococcus.  They  take  the  date  and  authority  of  the  author  who  first  used  them 
in  a  subgeneric  sense  and  not  the  date  and  authority  of  the  author  who  first  pro- 
posed them,  since  according  to  the  International  Code,  section  48,  b,  the  generic 
and  specific  names  of  larval  cestodes  (Cysticercus,  Cocnurus,  Echinococcus,  Finna, 
Hydatigena,  Hydatis.  etc.)  are  not  entitled  to  the  law  of  priority. 

The  genus  Tainia  is  frequently  divided  into  three  subgenera  :  Cysticercus  (  =  Cys- 
totcenia\  Ccenurus,  and  Echinococcus.  Tasnia  must  be  substituted  for  the  first  sub- 
genus  (type  T.  aolium)  and  Echinococcifer  must  probably  be  substituted  for  the  third 
subgenus. 


214  TAPEWORMS  OF  HAKES  AND  RABBITS—  STILES. 


Egg  with  thin  transparent  shells;  frequently  in  egg  capsules;  in  some  cases 
scattered  through  the  segments;  head  nearly  always  armed  with  hooklets  on 
rostelluni;  larval  stage  a  cysticercoid ;  adults  in  birds  and  mammals. 

Dipylidiince  (p.  193),  27 
Anoplocephalince.1 

5.  Ova  with  well  developed  pyriform  body;   genital  canals  pass  dorsally  of  longi- 

tudinal canals  and  main  nerve  trunk  (not  yet  satisfactorily  demonstrated  for 
Andrya) 6 

Ova  without  pyriform  body;  genital  canals  pass  dorsally  of  ventral  canal  and  nerve, 
but  ventrally  of  dorsal  canal  (not  yet  demonstrated  for  Stilesia  centripunctata) .  10 

6.  Uterus  a  transverse  tube,  generally  continuous,  in   some  of  the  double-pored 

forms  divided  (double),  one  tube  for  each  ovary,  and  generally  provided  with 
proximal  and  distal  pouches;  dorsal  canal  lies  dorsal  to  dorso-lateral  of  ventral 

canals ;  pedunculated  prostatic  gland  absent 7 

Uterus  more  like  a  network  in  its  early  stages;  afterwards  the  boundaries  of  the 
meshes  nearly  or  quite  disappear  and  the  uterus  (or  uteri)  appears  like  a  sac. .  9 


1  The  genera  of  Anoplocephalince  may  also  be  determined  by  the  following  key,  which 
although  simpler  than  the  one  given  above,  is  much  more  artificial : 

Anoplocephalina. 

A.  Genital  pores  single B 

Genital  pores  double F 

B.  Genital   pores   unilateral    or  irregularly   alternate  with   decided   tendency  to 

unilaterality .• C 

Genital  pores  regularly  alternate  or  nearly  so;  pedunculate  prostatic  glands 
absent  D 

C.  Pores  unilateral;  testicles  unilateral  or  nearly  so  in  median  field  opposite  pores; 

uterus  transverse  with  proximal  and  distal  pouches;   pedunculate  prostatic 

gland  absent Anoplocephala,  1 1 

Pores  irregularly  alternate  with  decided  tendency  to  unilaterality;  uterus  forms 
a  network  in  its  younger  stage,  later  the  meshes  become  nearly  or  quite 
obliterated;  testicles  extend  across  the  median  field  to  or  beyond  the  ovary; 
distinct  pedunculated  round  or  elongate  prostatic  gland  present. ..  Andrya,  26 

D.  Egg  without  pyriform  body E. 

Egg  with  well-developed  pyriform  body;   uterus  transverse  with  thin- walled 

proximal  and  distal  pouches;  genital  canals  pass  dorsally  of  dorsal  and  ventral 
canals  and  main  longitudinal  nerve  trunk Bertia,  14 

E.  Uterus  single  and  transverse,  without  pouches,  or  double,  a  cornucopia-like  egg  pouch 

being  present  near  each  ventral  canal,  the  median  transverse  connecting  canal  not 
visible;  genital  canals  pass  dorsally  of  nerve  and  ventral  canal,  but  ventrally  of  dor- 
sal canal  (not  shown  for  S.  centripunctata) *. Stilesia. 

Uterus  transverse  but  undulate,  with  thick-walled  asconspore  or  cornucopia-like  egg 
pouches;  genital  canals  pass  dorsally  of  nerve  and  ventral  canal,  but  ventrally  of 
dorsal  canal Thysanosorna. 

F.  Egg  without  pyriform  body;  uterus  transverse  and  undulate,  with  thick-walled  ascon- 

spore or  cornucopia-like  egg  pouches;  genital  canals  pass  dorsally  of  ventral  canals 

and  nerves,  but  ventrally  of  dorsaj  canal Thusanosoma. 

Egg  with  well- developed  pyriform  body ;  genital  canals  dorsally  of  ventral  an  I 
dorsal  canals  and  nerves G 

G.  Uterus  a  transverse  tube,  single  or  double,  with  thin-walled  egg  pouches ;  dorsal 

canal  dorsal  to  dorso-lateral  of  ventral  canal ;  vagina  ventral  of  cirrus  pouch 

on  both  sides  of  the  segment Cittotcenia,  19 

Uterus  reticulate,  double;  dorsal  canal  dorsal  to  dorso-median  of  ventral  canal;  vagina 
ventral  of  right  cirrus  pouch,  dorsal  of  left  cirrus  pouch Moniezia. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  215 

7.  Genital  pores  single 8 

Genital  pores  double;  vagina  ventral  of  cirrus  pouch  on  both  sides  of  the  seg- 
ment     Cittotcenia  (p.  170),  19 

8.  Pores  unilateral;  testicles  unilateral,  or  nearly  so,  in  median  field. 

Anoplocephala  (p.  150),  11 

Pores  regularly  or  irregularly  alternate;  testicles  extend  ncrosg  the  median  field 
(not  shown  as  yet  for  B.  studeri  and  B.  satyri) Bertia  (p.  160),  14 

9.  Genital  pores  irregularly  alternate  with  marked    tendency  to  unilaterality ; 

pedunculated  round  or  elongate  prostatic  gland  near  ventral  canal  on  pore  side 
of  segment;  dorsal  canal  lies  dorsal  to  dorso  (?  lateral)  of  ventral  canal. 

Andrya  (p.  154),  26 

Genital  pores  double;  pedunculated  prostatic  gland  absent;  vagina  ventral  of  right 
cirrus  and  dorsal  of  left  cirrus Moniezia.1 

10.  Uterus  single,  transverse,  but  undulate,  with  asconspore  or  cornucopia-like  egg  pouches; 

genital  pores  single  or  double;  testicles  form  a  band  in  median  field  or  are  divided 

into  two  lateral  groups,  one  in  each  lateral  field Thysanosoma.* 

Uterus  single  and  transverse,  or  double,  being  represented  by  a  cornucopia-UTce pouch 
on  each  side  of  the  segment;  genital  pores  irregularly  alternate;  testicles  in  two  lat- 
era  I  groups  and  absent  from  median  line Stilesia.* 

Anoplocephala. 

11.  Head  with  posterior  lobes;  body  may  attain  8  to  25  mm.  long  by  3  to  15  mm.  wide; 

head  2  to  3  mm.  broad.     Host:  Horse  (Equus  caballus) A.perfoliata. 

Head  without  posterior  lobes 12 

12.  Head  4  to  6  mm.  broad;  strobila  9  to  80  cm.  long  by  5  to  20  mm.  broad.   Host:  Horse 

(Equus  caballus)  and  Mule  (Equus  asinus  X  caballus) A.plicata.4 

Head  less  than  1  mm.  broad 13 

13.  Strobila  10  mrn.  long  by  1.5  to  2.25  mm.  broad;   contains  10  to  28  segments; 

distal  segments  may  become  nearly  as  long  as  broad;  genital  glands  confined 
to  proximal  third  of  worm ;  testicles  15  to  30  111  each  segment ;  cirrus  pouch 
0.48  mm.  long.  Host:  European  rabbit  (Lepus  cuniculus)  and  Mountain  hare 
(L.  variabilis) A.  wimerosa. 

Strobila  attains  6  to  30  mm.  long  by  4  to  6mm.  broad;  35  to  53  segments  present,  the 
distal  4  to  8  completely  filled  with  ova;  testicles  60  to  100  in  each  segment;  cirrus 
pouch  attains  0.8  mm.  in  length.  Host:  Horse  (Equus  caballus). 

A.mamillana  (p.  153)' 

Strobila  10  to  16  cm.  long  by  6  to  8  mm.  broad;  60  to  80  testicles  present.  Host: 
Arctomys,  sp A.  transversaria  (p.  154). 


1 1891,  Moniezia,  R.  BLANCHARD,  Me"m.  Soc.  zool.  France,  IV,  p.  187.  Type,  Twnia 
expansa,  Rudolphi,  1810. 

For  the  species  of  this  genus  see  Stiles  &  Hassall,  1893,  A  Revision  of  the  Adult 
Cestodes  of  Cattle,  Sheep,  and  Allied  Animals,  Bull.  4,  Bureau  of  Animal  Industry, 
U.  S.  Dept.  Agriculture. 

>21835,  Thysanosoma,  DIESING,  Med.  Jahrb.  d.  ost.  Staat.  n.  Folge,  VII,  pp.  105-111, 
pi.  in.  Type,  Thysanosoma  actinioides,  Diesing,  1835. 

For  the  species  of  this  genus  see  Stiles  in  Stiles  &  Hassall,  1893,  Bull.  4,  Bureau 
of  Animal  Industry. 

3  1893,  Stilesia,  RAILLIET,  Traite  de  Zool.  me'd.'et  agric.,  I,  pp.  277-278.  Type,  Tcenia 
globipunctata,  Rivolta,  1874. 

For  the  species  of  this  genus  see  Stiles  in  Stiles  &  Hassall,  1893,  Bull.  4,  Bureau 
of  Animal  Industry. 

4Doubtful  whether  this  specific  term  should  stand,  as  it  is  antedated  by  equina, 
1781,  and  magna,  1789. 


216  TAPE  WORMS  OF  HAEES  AND  EA  BEITS- STILES. 


Inertia. 

14.  Hosts:  Rodents;1  cirrus  pouch  distinctly  muscular,  prominent,   with  vesicula 

seminalis;  receptaculum  seminis  round  and  prominent;  pores  very  regularly 

alternate ;  calcareous  corpuscles  present  or  absent 15 

Hosts:  Primates  and  lemurs;1  cirrus  pouch? ;  calcareous  corpuscles  present 16 

15.  Genital  pores  in  posterior  half  of  lateral  margin;  cirrus  pouch  about  0.5  mm.  long; 

posterior  segments  show  a  tendency  to  Itecome  narrow  and  thick;  strobila  attains  33 
mm.  in  length  by  6  mm.  in  breadth,  and  contains  about  90  segments,  calcareous  cor- 
puscles not  observed.  Host:  Yellow-haired  porcupine  (Erethizoii  epixanthus)  and 

Canada  porcupine  (E.  dorsatus),  North  America IS.  americana  (p.  165) 

Genital  pores  in  about  the  middle  of  the  lateral  margin  ;  cirrus  pouch  about  0.4  mm. 
long ;  posterior  segments  do  not  become  narrow ;  strobila  attains  23  to  47  mm.  in 
length  by  5  to  6.5  mm.  in  breadth,  and  contains  60  to  95  segments;  calcareous 
corpuscles  present.  Host:  Rabbit  (Lepus,  sp.  ?),  North  America. 

B.  americanaleporis  (p.  167). 

16.  Cirrus  pouch  not  so  prominent  as  vagina,  and  with  but  weak  musculature;  receptacu- 

lum seminis  round  to  elongate  and  not  very  prominent;  pores  irregularly  alternate; 
ovary  confined  to  pore  side  of  median  field;  uterus  transverse  with  proximal  and  distal 
pouches 17 

Cirrus  pouch  very  muscular;  receptaculum  seminis  elongate;  pores  irregularly  alter- 
nate; ovary  extends  to  aporose  portion  of  median  field;  uterus  transverse  with  prox- 
imal and  distal  pouches;  strobila  24  to  220mm.  long  by  5  to  11  mm.  broad,  and  con- 
tains SO  to  400  segments.  Host:  Fining  lemur  (Galeopithecus  volans),  India. 

B.  plattica  (p.  164). 

Cirrus  pouch?  vesicula  seminalis?;  uterus  composed  of  30  to  35  polyhedral  packages.  18 

17.  Strobila  attains  140  mm.  or  more  in  length  by  8  to  10  mm.  in  breadth.     Host:  Black 

howler  ( Alouatta  caraya),  Paraguay  B.  mucronafa  (p.  162). 

Strobila  attains  84  mm.  or  more  in  length  by  6.5  mm  in  breadth.  Host:  Bonnet  mon- 
key (Macacus  siuicus),  India B.  conferta  (p.  163). 

18.  Strobila  attains  130  mm.  in  length  by  15  mm.  in  breadth  by  2.5 mm.  in  thickness;  geni- 

tal pore  alternates  very  regularly;  ova  53  to  60  //.     Host:  Chimpanzee  (Aiithropo- 

pithecus  troglodytes) B.  studeri  (p.  161). 

Strobila  attains  245  mm.  or  more  in  length  by  10  mm.  in  breadth  by  2  mm.  in  thick- 
ness; pores  irregularly  alternate;  cirrus  pouch  claviform;  ova  35  to  38  /<  by  30  to 
32ju.  Host:  Oran-utau .  (Simia  satyrus) B.  satyri  (p.  161). 

Cittotainia. 

19.  Cirrus  pouch  muscular,  generally  pyriform,  with  distinct  outline,  and  swollen 

proximally  by  the  vesicula  serainalis Marmota>-Group,  20 

Cirrus  pouch  tubular,  resembling  the  nozzle  of  a  hose,  of  equal  or  nearly 
equal  diameter  throughout  its  entire  length,  and  not  swollen  proximally  by 
any  prominent  vesicula  seminalis Pectinata-Group,  23 

20.  Testicles  arranged  in  a  band  extending  across  the  median  field 21 

Testicles  comparatively  few  in  number,  absent  from  median  line,  arranged  in  two 

triangular  groups,  one  corresponding  to  each  ovary;  cirrus  pouch  small, 
0.16  mm.  long,  scarcely  reaching  the  longitudinal  nerve;  pores  in  distal  part 
of  lateral  margin;  female  glands  near  ventral  canal;  dorsal  canal  dorsal  or 
dorso-lateral  of  ventral  canal;  strobila  attains  80  cm.  in  length  by  10  mm.  in 
breadth.  Host:  Common  wild  and  tame  rabbits  (Lepus  cuniculus  and  L.  cnui- 
culus  domesticus),  Europe C.  ctenoides  (p.  179). 

21.  Testicles  not  confined  to  space  between  the  ovaries 22 

Testicles  confined  to  distal  portion  of  segment  between  oraries;  cirrus  pouch  about  0.5 

mm.  long  by  0.17  mm.  broad,  crosses  the  canals  ;  female  glands  some  distance  from 


1  Characters  satisfactory  for  an  analytical  key  are  still  lacking. 


NO.  1105.  PROCEEDINGS  OF  THE  NA  TIONAL  MUSE UM.  217 

lateral  canals  ;  transverse  uterus  single  with  proximal  and  distal  pouches;  dorsal 
canal  'between,  ventral  canal  and  nerve;  stroblla  may  attain  112  mm.  long  by  5  to  IS 
mm.  broad.  Host :  Marmot  ( Arctomys  inarmota),  Europe.  C.  marmotm  (  p.  172). 

22.  Cirrus  pouch  -very  large  and  prominent,  may  attain  1.12  mm.  long  by  0.32  mm. 

broad;  pores  at  or  near  distal  corner  of  lateral  margin;  female  glands  not  far 
removed  from  dorsal  canal;  several  accessory  ventral  canals;  strobila  may 
attain  80  cm.  long  by  15mm.  broad.  Host:  Wild  rabbit  (Lepus  cuniculus), 

Europe C.  denticulata  (p.  174). 

Cirrus  pouch  small,  0.24  mm.  long;  female  glands  considerably  removed  from  dorsal 
canal;  dorsal  canal  lateral  or  dorso-lateral  of  ventral  canal ;  reccptaculum  seminis 
very  large;  segments  imbricate;  strobila  attains  40  mm.  long  by  5.5  mm.  broad. 
Host:  Prairie  gopher  (Geornys  bursarius),  North  America..  C.  prwcoquis  (p.  181). 

23.  Cirrus  pouch  about  1  mm.  long,  extends  some  distance  median  of  longitudinal 

canals ;  testicles  arranged  in  a  band  in  distal  portion  of  median  field,  extend- 
ing beyond  ovaries  from  canal  to  canal;  ovary  some  distance  from  longitudi- 
nal canals;  uterus  single,  with  well-developed  proximal  and  distal  pouches; 
strobila  attains  40  cm.  in  length  by  8  to  10  mm.  in  breadth.  Hosts:  Common 
hare  (Lepus  timidus)  and  mountain  hare  (L.  variabilis)  Europe. 

C.pectinata  (p.  184). 

Cirrus  pouch  not  over  0.5  mm.  long,  extends  scarcely  median  of  lateral  canals'; 
uterus  single  or  double  in  the  same  strobila.  Hosts :  Lepus,  North  America.  24 

24.  Testicles  in  two  groups,  one  for  each  ovary,  extending  laterally  of  ovaries  but 

absent  from  median  field  ;  strobila  attains  57  mm.  or  more  in  length  by  10  mm. 

in  breadth.     Host:  Cottontail  rabbit  (L.  sylraticus) C.  perplexa  (p.  189). 

Testicles  in  a  band  confined  to  distal  portion  of  median  field  between  the 
ovaries;  strobila  attains  100  to  180  mm.  in  length  by  10  mm.  in  breadth. 
Host:  Cottontail  rabbit  (L.  sylvaticus)  and  marsh  hare  (L.  palustris). 

C.  variabilis  (p.  190),  25. 

25.  Strobila  nearly  always  more  than  3  mm.  broad ;   posterior  flap   of  segments 

nearly  straight;  segments  slightly  imbricate:  genital  pore  in  about  the 
middle  of  the  lateral  margin.  Host:  Cottontail  rabbit  (L.  sylvaticus). 

C.  variabilis  (p.  192). 

Strobila  nearly  always  more  than  3  mm.  broad;  posterior  flap  of  segments 
lobed;  segments  imbricate;  genital  pores  generally  in  distal  half  of  lateral 

margin.     Host :   Marsh  hare  (L.  palustris) C.  variabilis  imbricata  (p.  193). 

Strobila  rarely  over  2  mm.  broad;  posterior  flap  straight;  segments  not  imbri- 
cate; genital  pores  generally  in  distal  portion  of  lateral  margin,  generally 
double,  occasionally  single.  Host:  Cottontail  rabbit  (L.  sylvaticus'). 

C.  variabilis  angusta  (p.  193). 
Andrya  (European). 

26.  Genital  pores  near  posterior  corner  of  segment ;  prostatic  gland  elongate ;  head 

about  1  mm.  in  diameter;  testicles  comparatively  few  in  number,  confined 
chiefly  to  aporose  side  of  median  field;  cirrus  pouch  0.32  to  0.34  mm.  long; 
strobila  attains  60  to  80  cm.  in  length  by  5  mm.  in  breadth.  Host:  Common 

hare  ( Lepus  timidus),  Europe - A .  rhopalocephala  (p.  155). 

Genital  pores  in  about  the  middle,  or  in  the  distal  half  of  the  lateral  margin; 
prostatic  gland  round;  head  about  0.5  mm.  in  diameter;  testicles  about  50  in 
number,  scattered  through  entire  median  field;  cirrus  pouch  0.4  to  0.48  mm. 
long;  strobila  attains  100  cm.  in  length  by  8  mm.  in  breadth.  Host:  Wild 
rabbit  (Lepus  cuniculus),  Europe -  ^-  cuniculi  (p.  158). 

DipylidiincB. 

27.  Suckers  unarmed  — -  •  ** 

Suckers  armed  with  minute  booklets  (with  four  exceptions  known  only  in 

birds) 3 

28.  Genital  pores  double ^ 29 

Genital  pores  single,  irregularly  alternate  or  unilateral 31 


218  TAPE  WORMS  OF  HARES  AND  RAB  BITS— STILES.  VOL.  xix. 

29.  Two  submedian  ovaries  in  each  segment 30 

One  median  ovary  in  each  segment Amabilia.1 

30.  Several  rows  of  hooks  upon  rostellum Dipylidium.2 

A  single  row  of  hooks  upon  rosteUum Cotugnia.* 

31.  Dorsal  root  of  hooks  much  longer  than  ventral  root  or  prong ;  ventral  root  very  short; 

hooks  8  to  12  (rarely  to  26}  in  number;  known  only  in  birds Drepanidotwnia.* 

Dorsal  root  of  hooks  about  the  same  length  as  ventral  root  and  prong 32 

32.  Dorsal  root  shorter  than  prong  or  ventral  root ;  hooks  generally  less  than  20  in  number 

(range  from  10  to  26}  ;  genital  pores  unilateral  or  irregularly  alternate  ;  known  only 

in  birds Dicranolamia.5 

Dorsal  root  longer  than  prong  or  ventral  root ;  if  rosteUum  is  armed,  there  are  24  to  30 
hooks  present ;  genital  pores  unilateral  (on  left  of  segments)  ;  three  testicles  nor- 
mally present  in  each  segment ;  retractile  rostellum  armed  with  minute  hooklets  or 
rudimentary  and  unarmed;  eggs  with  three  envelopes  ;  parasitic  in  man,  chiroptera, 
insectivora,  rodents,  and  insectivorous  birds  ;  larval  stage  in  insects  or  myriapoda.  - 

Hymenolepis.* 

33.  Hooks  on  suckers  arranged  in  circular  rows  on  border;  hooks  on  rostellum  resem- 

ble a  hammer  in  form  and  are  arranged  in  a  double  row;  with  four  exceptions 
known  only  in  birds Davainea  (p.  194),  35. 


'1893,  Amabilia,  DIAMARE,  Boll.  Soc.  Nat.  Napoli,  1.  ser.,  VII  (1-2),  p.  13.     August 

28,  1893.     Type,  Tcenia  lamelligera,  Owen,  1835. 
?  1896,  Diploposthe,  JACOBI,  Zool.  Auzeiger,  XIX,  No.  505,  June  15,  1896,  pp.  268-269. 

Type,  Tcenia  Icvvis,  Diesiug. 

See  Stiles,  1896,  Bull.  12,  Bureau  of  Animal  Industry,  p.  31.     From  the  diagnosis 
given  by  Jacobi  I  fail  to  see  how  Diploposthe  diifers  from  Amabilia. 
£1863,    Dipylidium,  R.  LEUCKART,   Die  menschlicheu  Parasiten,  I,  p.  400.      Type, 

Tcenia  elfyrtica,  Batsch,  1786  =  Tcvnia  canina,  Linnaeus,  1758. 
1858,   "  Alyselminthus,    ZEDER,   1800    pp.,"    of  Weinland   with    Tcvnia    cucumerina, 

Bloch,  1782  —  Tamia  canina,  Liumeus,  1758,  as  type. 
1884,   Micro  tcenia,  SEDGWICK,  as  subgenus  in   CLAUS    &   SEDGWICK,    Elementary 

Text-book  of  Zoology,  I,  p.  336.      Type,  Tamia  cucumerina,  Bloch,  1782  = 

Tcenia  canina,  Linnaeus,  1758. 

For  species  and  bibliography  see  Diamari's  monograph  (1893)  II  Gen  ere  Dipylidium, 
Lkt.,  Atti  R.  Accad.  Scienze  fisische  e  matematiche,  2  ser.,  VI,  No.  7,  31  pp.,  pis. 
I-III. 
3 1873,  Cotugnia,  DIAMARI,  Boll.  Soc.  Nat.  Napoli,  1.  ser.,  VII  (1-2),  p.  11.     August 

28,  1893.     Type,  Tfenia  digonopora,  Pasquale,  1890. 
See  Stiles,  1896,  Bull.  12,  Bureau  of  Animal  Industry,  p.  29. 
4 1892,  Drepanidotcenia,  RAILLIET,  Bull.  Soc.  zool.  France,  XVII,  p.  116.     Type,  Tcmia 

lanceolata,  Bloch,  1782. 
?  1896,  Choanotcenia,  RAILHET,  Rec.  me"d.  ve"t.,  8  ser.,  Ill  (5),  March  15,  1896,  p.  159. 

Type,  Tcenia  infundibuliformis,  Goeze,  1782. 

For  species  and  bibliography  of  this  genus  see  Stiles,  1896,  Bull.  12,  Bureau  of  Animal 
Industry,  pp.  36-45,  60-61. 
61892,  Dicranotosnia,  RAILLIET,  Bull.  Soc.  zool.  France,  XVII,  p.  116.     Type,  Tcenia 

coronula,  Dujardin,  1845. 

For  species  and  bibliography  of  this  genus  see  Stiles,  1896,  Bull.  12,  Bureau  of 
Animal  Industry,  pp.  32-36. 
61858,  Hymenolepis,  WEINLAND,  Human  Cestoides,  p.  49,  52.      Type,  Tcenia  Jlavopunc- 

tata,  Weinland,  1858=  Toenia  diminuta,  Rudolphi,  1819. 
1858,  Lepidotrias,  WEINLAND,  as  subgenus  of  Hymenolepis,  Human  Cestoides,  p.  52. 

Type,  Tceniamurina,  Dujardin,  1845,  proposed  by  Weinland,  but  this  subgenus 

includes  the  type  of  the  genus. 

For  species  and  bibliography  of  the  genus  see  R.  Blanchard,  1891,  Hist.  zool.  et 
med.  des  Teiriades  du  genre  Hymenolepis,  Weinlaud.     Paris.     112  pp.,  22  figs. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  219 


Hooks  on  stickers  arranged  in  several  transverse  rows ;  hooks  of  infundibulum  very 

small  (4  /<)  arranged  in  a  single  row  ;  known  only  in  birds Ophryocotyle.1 

Hooks  on  suckers  arranged  in  one  median  set  and  two  lateral  sets  ;  hooks  on  rostellum 
provided  with  long  dorsal  root  and  arranged  in  a  single  row  ;  known  only  in  birds. 

Ecliinocotyle* 
Subfamily  f 

34.  Anterior  extremity  of  strobila  expanded  in  form  of  a  hammer Fimbriaria.* 

A  n  terior  segments  become  calci/orm  and  function  as  pseudoscolex Idiogenes.4 

Davainea. 

35.  Found  in  birds  ;  see  Stiles,  1896,  5  p.  45. 

Found  in  mammals r 36 

36.  Genital  pores  unilateral ;  a  single  egg  in  each  egg  capsule 37 

Genital  pores  generally  alternate;  strobila  attains  85  mm.  or  more  in  length  by 

3  mm.  in  breadth;  suckers  not  invaginated;  eggs  grouped  3  to  15  in  each  egg 
capsule.  Host:  Eastern  Jackass  hare  (Lepus  melanotis)  and  Cottontail  rabbit 
(L.  sylvaticus) I),  salmoni  (p.  198). 

37.  Suckers  (always?)  invaginated;  strobila  attains  105mm.  long  by  3  mm.  broad. 

Host :  Arizona  cottontail  (L.  arizonm) D.  retractilis  (p.  195). 

Suckers  not  invaginated 38 

38.  Rostellum  armed  with  double  row  of  about  90  hooks,  IS  /.i  long  ;  strolnln  attains  250 

to  300  mm.  long,  composed  of  500  to  600  segments.     Host :  Man  (Homo  sapiens). 

D.  madagascariensis  (p.  194). 

Rostellum  large,  armed  with  (?)  a  single  row  of  numerous  minute  hooklets ;  suckers 
armed  with  8  to  10  rows  of  hooks.  Host:  Common  Indian  Pangolin  (Manis  peu- 
tadactyla) D.  contorta  (p.  195). 

COMPENDIUM   OF   THE   PARASITES   ARRANGED   ACCORDING   TO   THEIR 

HOSTS. 

In  the  following  list  are  given  the  hosts  of  the  parasites  discussed  in 
this  paper,  so  far  as  they  are  known  to  me.  I  have  personally  exam- 
ined the  species  starred  (*)  for  the  hosts  under  which  they  are  given. 
For  the  name  of  the  collector  in  each  case  see  text.  The  numbers  given 
with  the  names  of  the  hosts  are  those  of  von  Linstow's  Compendium 
and  Nachtrag.  The  geographical  distribution  refers  to  the  host.  One 


'1870,  Ophryocotyle,  FRIIS,  Videnskab.  Meddel.  fra  den  Naturhist.    Foren.  Kjoben- 

havn  [aar  1869],  1870,  pp.  121-124,  pi.  I.     Type,  Ophryocotyle  proteus,  Friis, 

1870. 

R.  Blaiichard  gives  the  date  as  1869,  Braun  gives  it  1870.     Original  not  at  my  dis- 
posal.    For  species  see  Stiles,  1896,  Bull.  12,  Bureau  of  Animal  Industry,  p.  56. 
2 1891,  Echinocotyle,  R.  BLANCHARD,  Bull.  Soc.  zool.    France,    IV,  p.  423.     Type,  E. 

rosseteri,  R.  Blan chard,  1891. 
31802,  Fimbriaria,   FROLICH,  Der  Naturforscher,    XXIX,    pp.    13-14.     Type,    Tanta 

malleus,  Goeze,  1782. 
1850,  Ithynchotwnia,  DIESING  (as  section  or  subgenus  of  Tojnia},   Systema  Helmiii- 

thum,  I,  p.  521.     Type,  Twnia  malleus,  Goeze,  1782. 
1892,  Epision,  LINTON,  Proc.  U.  S.  Nat.  Mus.,  XV,  p.  100.     Type,  Epision  plicatus, 

Linton,  1892. 
4 1868,  Idiogenes,  KRABBE,  Videns.  Meddel.  fra  d.  Naturhist.  Foren.  Kjobeuhavn  (for 

Aaret  1867),  1868,  pp.  122-126.     Type,  Idiogenes  otidis,  KRABBE,  1868. 
6  Stiles,  1896,  Report  upon  the  Present  Knowledge  of  the  Tapeworms  of  Poultry, 

Bull.  12,  Bureau  of  Animal  Industry,  U.  S.  Department  of  Agriculture. 


220  TAPEWORMS  OF  HARES  AND  RABBITS—  STILES.  VOL.XIX. 


or  two  parasites  not  discussed  in  this  paper  are  included  in  the  com- 
pendium for  reasons  obvious  in  each  case. 

1  have  elsewhere  (1896)  made  a  plea  for  the  adoption  of  a  modern  sys- 
tem of  nomenclature  for  the  hosts,  and  have  endeavored  to  introduce 
here  the  correct  names  for  most  of  the  hosts  cited.  In  the  main,  Flower 
and  Lydekker's  Mammals  Living  and  Extinct  (1891)  has  been  followed; 
in  the  few  cases  that  I  have  departed  from  the  names  given  in  that  work 
I  have  done  so  upon  the  advice  of  Dr.  T.  S.  Palmer.  In  order  to  pre- 
vent confusion  by  this  change  in  host  names.,  I  have  added  the  names 
used  by  von  Linstow  in  his  compendium  or  by  other  authors  in  their 
writings,  cross-referencing  the  synonyms  to  the  proper  name.  It  is 
impossible  to  give  a  monographic  list  of  the  genus  Lepus  and  to  estab- 
lish the  correct  names  of  all  the  forms  until  that  genus  is  revised,  by  a 
specialist  in  mammalogy. 
ALOUATTA  CARAYA  (Humboldt,  1811).  Black  Howler. 

(Stentor  caraya,  Humboldt;  20.  Mycetes  niger ;  31.   Celms  caraya,  Fischer). 

(Southern  Brazil,  Paraguay,  Bolivia.) 

*  Bertia  mucronata. 
Tcenia  megastoma. 

ANTHROPOPITHECUS  TROGLODYTES  (Linnaeus,  1758).     Chimpanzee. 
(2a.  Simia  troglodytes ;   Troglodytes  niger  Geoff.) 
(Western  and  Central  Equatorial  Africa.) 
Bertia  studeri. 
92.  ARCTOMYS  MARMOTA  (Linnaeus,  1758).     Alpine  Marmot. 

(Mountains  of  southern  Europe — Alps,  Pyrenees,  and  Carpathians.) 
*  CittotoRnia  marmottv. 
926.  ARCTOMYS,  sp.  ? 

(Turkestan.) 

Anoplocephala  transversaria. 

31.  CEBUS  CARAYA,  Fischer  vide  ALOUATTA  CARAYA. 
30.  CEBUS  CAPUCINUS  (Linnaeus,  1758).     Weeping  Capnchin. 
(Paraguay  to  United  States  of  Colombia.) 
"Taenia  No.  2,  Gottheil,"  p.  164. 
248.  EQUUS  CABALLUS  (Linnaeus,  1758).     Horse. 
*AnoplocepJiala  mamillana. 
Anoplocephala  perfoliata. 
Anoplocephala  plicata. 
Anoplocephala  plicata  pediculata. 
Anoplocephala  plicata  strangulata. 
124.  ERETHIZON  DORSATUS  (Linnaeus,  1758).     Canada  Porcupine. 

(Boreal  region  of  eastern  North  America.) 
*  Bertia  americana. 
Tosnia  laticephala,  Leidy. 

?  "  Tcenia pectinata"  with  unilateral  pores=?,  recorded  by  Cobbold,  1862. 
ERETHIZON  EPIXANTHUS,  Brant.     Yellow-haired  Porcupine. 
(Mountains  of  Western  United  States.) 

*  Bertia  americana. 

GALEOPITHECUS  VOLANS  (Linnaeus,  1758).    Flying  Lemur,  Common  Colngo  or  Cobego. 

(Sumatra,  Borneo,  Java,  Malay  Peninsula,  Tenasserim,  and  Siain.) 
Bertia  plastica. 
GEOMYS  BURSARIUS  (Shaw,  1800).     Prairie  Gopher. 

(Upper   Mississippi  Valley,  southward  to   southern  Illinois,  Missouri,  and 
eastern  Kansas.) 

*  Cittotcenia  prcecoquis. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  221 


1.  HOMO  SAPIENS,  Linmeus,  1758.     Man. 

BothriocepUalus  cordatns. 

*  Bothrloceplialus  latns. 

Davainea  madagascariensis 

Dipylidium  caninum. 

Hymcnolepis  diminuta. 

Hymenolepis  murina. 

Krabbea  grandis* 

*Tcenia  confasa. 

*Tcenia  saginata. 

*Tamia  solium. 

12.  INUUS  CYNOMOLGUS  vide  MACACUS  CYNOMOLGUS. 
LEPUS  AMERICANUS,  Erxleben,  1777.     Northern  Hare. 

(Wooded  districts,  New  England  to  Minnesota,  and  south  to  Virginia,  along 

the  Alleghanies. ) 

?  "  Toenia pectinata,"  reported  by  Curtice, '  1892,  p.  232. 
LEPUS  ARIZONA,  Allen,  1877.     Arizona  Cottontail. 

(Arizona,  southern  Nevada,  and  desert  region  of  southern  California.) 

*  Davainea  retractilis. 

137.  LEPUS  CUNICULUS.    Common  European  Wild  Rabbit. 

Andrya  cuniculi.    See  p.  155. 
* Anoplocephala  wimerosa. 
*Cittotamia  ctenoides. 
*Cittota'nia  denticulata. 

137a.  LEPUS  CUNICULUS  DOMESTICUS.    Common  Domesticated  Rabbit. 
*Cittot(vnia  ctenoides. 

?  "  Tcema  pectinata,"  *  reported  by  Curtice,  1892,  p.  232. 
LEPUS  MELANOTIS,  Mearns,  1890.     Eastern  Jackass  Hare. 

(Plains  from  Texas  to  Nebraska,  west  to  Rocky  Mountains.) 

*  Davainea  salmoni. 

LEPUS  PALUSTRIS,  Bachrnan,  1837.     Marsh  Hare. 

(Marshy  lowlands  of  South  Atlantic  States,  from  North  Carolina  southward.) 
*Cittot(vnia  variabilis  imbricata. 

138.  LEPUS  SYLVATICUS,  Bachman,  1837.    Cottontail. 

(Eastern  United  States.) 
*CUtot(Knia  perplexa. 
*CitlotcKnia  variabilis. 
*Cittotce/nia  variabilis  angusta. 
^Davainea  salmoni. 
LEPUS  TEXIANUS,  Waterhouse,  1848.     Black-Tailed  Jack-Rabbit. 

(Great  Basin. ) 

"An  undescribed  Tamia,"2  reported  by  Curtice,  1892,  p.  233. 
140.  LEPUS  TIMIDUS,  Linnajus,  1758.     Common  European  Hare. 

(All  parts  of  Europe  except  the  north  of  Russia,  the  Scandinavian 

peninsula,  and  Ireland.) 
?  Andrya  cuniculi.     See  p.  155. 

*  Andrya  rhopalocephala. 

*  Cittotamia  pectinata. 

139.  LEPUS  VARiAiiiLis,  Pallas,  1778.    Mountain  Hare. 

(Northern  Eurasia.) 
* AnoplocepJiala  ivimerosa. 
*Cittotcenia  pectinata. 

ll  am  unable  to  trace  this  specimen,  but  most  of  Curtice's  T.  pectinata  is  C.  variabilis. 
-Impossible  to  tell  definitely  what  species  Curtice  referred  to,  but  possibly  the 
parasite  is  Davainea  salmoni  and  the  host  L.  melanotis. 


222 


TAPEWORMS  OF  HARES  AND  RABBITS— STILES. 


LEPUS,  SP.  ? 

(Some  North  American  species.) 
*Bertia  americana  leporis. 
?LEPUS  SP.?  (?WASHINGTONI,  S.  F.  Baird,  1855).     Western  Varying  Hare. 

(Puget  Sound.) 
* Cittotwnia  (?)  varidbilis. 

MACACUS  CYNOMOLGUS  (Linnaeus,  1758).     Crab-eating  Macaque. 
(12.  Innus  eynomoh/us.) 

(Malay  Peninsiila  and  Philippine  Archipelago.) 
"  Tcenia  No.  1,  Gottheil."    See  p.  1S4. 
"  Tcenia  No.  2,  Gottheil."     See  p.  164. 
MACACUS  sixicus  (Linnaeus,  1771).     Bonnet  Monkey. 

(Southern  India.) 
Bertia  conferta. 
MAXIS  PEXTADACTYLA.     Common  Indian  Pangolin. 

(India  aud  Ceylon.) 
Davainea  contorta. 
20.  MYCETES  NIGER,  vide  ALOUATTA  CARAYA.« 

3.    PlTHECUS  SATYRUS,  vide  SlMIA  SATYRUS. 

SIMIA  SATYRUS,  Linnaeus,  1758.     Orang  or  Oran-Utan. 

(3.  Pitfiecus  satyrus.) 

(Sumatra  and  Borneo.) 

Bertia  satyr  i. 

2a.  SIMIA  TROGLODYTES,  vide  ANTHROPOPITHECUS  TROGLODYTES. 
TROGLODYTES  NIGER,  vide  AXTHROPOPITIIECUS  TROGLODYTES. 

BIBLIOGRAPHY 

By  ALBERT  HASSALL,  M.  R.  C.  V.  S. 

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BAIRD,  W. 

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DE  BLAINVILLE. 

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BRAUN,  M. 

1894.—  Wiiriner:  Vermes.  Bronn'sKlas- 
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1895.— Id.  Bd.  IV,  Lief.  38-42,  pp.  1167- 
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1896.— Id.  Bd.  IV,  Lief.  43-44,  pp.  1247- 

1294,  Taf.  XL  VIII. 
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1824. — Icones  Helminth um  sy sterna  Ru- 
dolphii  Entozoologicum  illustrautes. 
Viennae.     fol.     XVIII  pis. 
COBBOLD,  T.  S. 

1862. — Remarks  on  Tcenia  pectinata.  In 
a  letter  from  Dr.  T.  Spencer  Cob- 
bold,  M.  D.,  F.  L.  S.,  London  (Eng- 
land), to  Professor  Lawson,  Queen's 
College,  Kingston.  The  Canadian 
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pp.  394-395. 
CREPLIN,  F.  C.  H. 

1842. — Endozoologische  Beitrage.  1. 
tiber  Tcenia  denticnlata  Rud.  und  T. 
expansa  Rud.  Arch,  fiir  Naturge- 
schichte.  Jahrg.  VIII,  Bd.  I,  pp. 
315-339. 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM. 


223 


CURTICE,  C. 

1888.  —Early  stages  in  the  life  of  Twnia 
pectinata.  Science,  N.  Y.,  March  23; 
also,  Texas  Live  Stock  Journal,  April 
14,  1888. 

1892. — Parasites,  being  a  list  of  those 
infesting  the  domesticated  animals 
and  man  in  the  United  States.  Jour- 
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Veterinary  Archives,  Vol.  XIII,  New 
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DAVAINE. 

1870. — Note  sur  une  nouvelle  espece 
de  Tsenia,  recueillie  a  Mayotte  (Co- 
mores)  ;  suivie  de  1'examen  micro- 
scopique  de  ce  Taenia,  par  le  Dr.  Da- 
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(1869)  1870.  M<Sm.,  pp.  233-240,  PL 
VI. 

DlESING.  K.  M. 

1850. — Systeina  Helminthum.     Vol.  I. 

Vindobonse.     679  pp. 
1854. — Uber    erne    Naturgemiisse    Ver- 

theilung  der  Cephalocotyleen.     Sitz. 

d.  k.  Akad.  d.  Wiss.  Wien,  Bd.  XIII, 

pp.  556-616. 
1864.— Revision    der  Cephalocotyleen. 

Abtheilung  Cyclocotyleen.     Sitz.  d. 

k.  Akad.  d.  Wiss.  Wien,  Bd.  XLIX, 

pp.  357-430. 

FROLICH.  „ 

1802. — Beitrage  zur  Naturgeschichte 
der  Eingeweidewiiriner.  Der  Natur- 
forscher,  St.  XXIX,  Halle,  pp.  5-96, 
Taf.  I-II. 

GOEZE,  J.  A.  E. 

1782. — Versuch  einer  Naturgeschichte 
der  Eingeweidewiirmer  thierischer 
Kofper.  Blankenburg,  471  pp.,  35  pis. 

GURLT,  E.  F. 
1831.— Lehrbuch    der     pathologischen 

Anatomie  der  Haus-Siiugethiere.     1. 

Theil.     Berlin.     399  pp. 
JAKOBI,  A. 

(1894).  —  Bandwiirmer       im       Hasen. 

Deutsche  Jager-Zeitung,  Bd.  XXIV, 

No.  7,  p.  98. 
KRABBE,  A. 

-Bidrag    til   Kundskab    oin    Fu- 

glenes     Bsendelorme.       Kgl.    Dansk 

Vidensk.  Selsk.  Skrifter,  5.  R.,Natur- 

vid.  og  math:  Afd.  8,  Bd.  VI,  1870,  pp. 

251-363,  Taf.  I-IX.     (Author's  extras 

published  1869.) 


KRABBE,  A. — Continued. 

(1879). — Cestodes  ges.  von  A.  P.  Fed- 
schenko  auf  seiner  Reise  in  Turkes- 
tan. Verb.  d.  k.  Ges.  d.  Frde.  d.  Na- 
tur.,  Anthropol.  und  Ethuographie, 
T.  XXXIV,  Moskau. 

LtJHE. 

1895  A. — Beitrage  zur  Kenntnis  des  Ros- 
telhims  und  der  Scolexmusculatur  der 
Tsenien.  Zool.  Anz.  No.  453,  XVII 
Jhg.,  pp.  279-282. 

1895  B. — Mitteilnngen  iiber  einige  we- 
nig  bekannte  bez.  neue  siidamerikani- 
scho  Tsenien  des  k.  k.  naturhistori- 
schen  Hof-Museums  in  Wien.  Arch, 
f.  Naturg.,  61,  Jhg.,  Bd.  I,  pp.  199-212, 
Taf.  XI. 

MARIGUES. 

1778. — Observation  sur  des  vers  Te"nia 
trouv<?s  dans  le  ventrc  de  quolques 
lapins  sauvages.  Observations  sur. 
la  Physique,  etc.  (Rozier),  Tom.  XII, 
Paris,  pp.  229-231,  PI.  II,  fig.  3. 

MEYNER,  R. 

1895.— Anatomie  und  Histologie  zweier 
neuer  Taenien-Arten  des  Subgeuus 
Bertia.  Tccnia  (Bertia)  mucronata  n. 
sp  und  Tcenla  (Bertia)  conferta  n.  sp. 
Ein  Beitrag  zur  Kenntuiss  der  Cesto- 
den.  Halle.  Inaug.-Diss.,  106  pp.,2 
plates;  also  Zeitschr.  f.  Naturwiss., 
Leipzig,  LXVIII  Bd.,  1895,  pp.  1-106, 
Taf.  I-II. 

MONIEZ,  R. 

1880.— Etudes  snr  les  Cestodes.  Bull, 
scientif.  du  Depart,  du  Nord,  2(!  s6r., 
3e  ami.,  pp.  240-246, 

NEUMANN,  L.  G. 

1887.—  Traite  des  maladies  parasitaires 
noii  microbiennes  des  auimaux  do- 
mestiques.  Paris.  673pp.,  306  figs. 

1892  A.— Trait6  des  Maladies,  etc.  2°  e"d. 
of  Neumann,  1887.  Paris.  767  pp., 
364  figs. 

1892  B.— Neumann's  Parasites  and  Par- 
asitic Diseases  of  Domesticated 
Animals.  800  pp.,  364  figs.  Transla- 
tion of  Neumann,  1892  A,  by  Flem- 
ing. London. 

PAGENSTECHER,  H.  A. 

1877. — Zur  Naturgeschichte  der  Cesto- 
den.  Zeitschr.  f.  wiss.  Zool.,  Bd. 
XXX,  pp.  171-193,  Taf.  X. 


224 


TAPEWORMS  OF  HARES  AND  RABBITS— STILES. 


PALLAS,  P.  S. 

1781. — Bemerkuugen  iiber  die  Band- 
wiirmer  in  Mensclien  und  Thieren. 
Neue  nordische  Beytriige,  etc.,  Bd.  I, 
St.  Petersb.  u.  Leipzig1,  pp.  39-112, 
PI.  II-III. 

PETERS,  W. 

1871. — Note  on  the  Tamia  from  the 
Rhinoceros  lately  described  by  Dr. 
Murie.  Proc.  Zool.  Soc.  London,  pp. 
146-147,  fig.  1-2. 

RAILLIET,  A. 

1890. — Une  nouvelle  affection  parasi- 
taire  du  lievre  et  du  lapin  de  ga- 
renne.  Rev.  d.  Sc.  nat.  appli.  Bullet, 
bimens.  de  la  Soc.  Nat.  d'Accliniat.  de 
France,  Tom.  XXXVII,  No.  8,  Paris, 
pp.  345-352. 

1893. — Trait6  de  zoologie  me'dicale  et 
agricole.  1.  fasc. 

1896. — Quelques  rectifications  ii  la  no- 
menclature des  parasites.  Rec.  d. 
mdd.  vet.,  viiie  aer.,  Tom.  Ill,  No.  5, 
pp.  157-161. 

RIEHM,  G. 

1881  A. —  [Untersuchungen  an  den 
Bandwiirmer  der  Hasen  und  Kanin- 
chen.]  Zeitschr.  f.  d.  ges.  Natur- 
wiss.  Halle.  Dritte  Folge,  Bd.  VI, 
p.  200. 

1881  B.— Studieu  an  Cestoden.  Zeit- 
schr. f.  d.  ges.  Naturwiss.,  Dritte 
Folge,  Bd.  VI,  pp.  545-610,  Taf.  V-VI  ; 
also  Iiiaug.  Diss.  Halle  a.  S.  66  pp., 
2  Taf. 

RUDOLPHI,  C.  A. 

1804. — Beinerkungen  aus  dem  Gebiete 

der    Naturgeschichte,    Medizin    und 

Thierarzneykunde    auf   einer    Reise 

durch  einen  Theil  von  Deutschland, 

Holland    und    Frankreich.      1.    Th. 

Berlin. 

1805.— Id.     2.  Th.     Berlin. 
1810. — Entozoorum  sive  vermium  intes- 

tinalium  historia  naturalis.     Vol.  II, 

Pt.  2.     Ainstehedami. 
1814. — Erster  Nachtrag  zu  ineiner  Na- 

turg.    d.    Eingeweidewiirmer.     Ges. 

Naturf.  Freunde   Berlin,  Mag.  f.  d. 

neust.  Entd.  i.  d.  ges.  Naturkde.,  VI. 

Jhg.,  Berlin,     pp.  83-113. 
1819. — Entozoorum  Synopsis.     Berlin. 

811  pp.,  Tab.  I-IH. 


SETTI,  E. 

1891.— Sulle   Tenie    dell'    Hyrax    dello 

scioa.     Atti  d.  Soc.  Ligustica  di  Sci- 

euze    Naturali,   Ann.  II,   Vol.11,    11 

pp.,  1  Tav.     Reprint. 
1893.— Elminti    delF    Eritrea    e    delle 

regioiii  limitrofe.     Atti  Soc.  Ligust. 

di  Sci.  Nat.,  Vol.  IV,  21  pp.    Reprint. 
STILES,  CH.  WAHDELL. 

1893.— Beinerkuugen  iiber  Parasiten— 

17:  Ueber  die   topographisclie  Aua- 

toniie    des     Gefasssystems     in     der 

Familie  Taaiiad.e.     Cent.  f.  Bakt.  u. 

Paras.,  XIII,  Nos.  14-15.  pp.  457-405, 

12  Abbild. 
1894  A. — Bemerkung  iiber  Parasiten — 

20:  Ueber  die  Erhaltung  von  Typeu. 

Centralbl.  fiir  Bakt.  u.  Paras.,  XV, 

Nos.  13-14,  pp.  477-480. 

1894  B.— Note surles  Parasites— 31 :  Uue 
phase  precoce  du  Te"nias  du  lapin. 
(Notice    prelimiiiaire. )      Bull.    Soc. 
Zool.  de  France,  Tome  XIX,  pp.  163- 
165;  Translation  in,  The  Veterinary 
Magazine,  Vol.  II,  1895,  pp.  32-33. 

1895  A. — Notes    on    parasites — 36:    A 
double-pored     Cestode     with     occa- 
sional single  pores.     Cent.  f.  Bakt. 
u.  Paras.,  1.  Abth.,  XVII,  Nos,  13-14, 
pp.  457-459, 1  fig;  alsoiu,  The  Veter- 
inary Magazine,  Vol.   II,    1895,   pp. 
222*225,  1  fig. 

1895  B. — Notes  on  parasites — 38:  Pre- 
liminary note  to  "A  revision  of  the 
adult  Leporine  Cestodes."    The  Vet- 
erinary Magazine,  Vol.  II,  pp.  341- 
346;    also    in    C.    R.    d.   stances  du 
troisieme  Congres  International   de 
Zoologie.    Leyde.   1896.  pp.  347-351. 

1896. — Report  upon  the  present  knowl- 
edge of  the  Tapeworms  of  poultr37, 
Bulletin  No.  12,  Bureau  of  Animal 
Industry,  U.  S.  Dept.  Agric.,  Wash- 
ington, D.  C.  pp.  1-79,  "Pis.  I-XXI. 
STILES,  CH.  WARDELL,  and  HASSALL, 
ALBERT. 

1893.— A  Revision  of  the  adult  Cestodes 
of  cattle,  sheep,  and  allied  animals. 
Bulletin  No.  4,  Bureau  of  Animal 
Industry,  U.  *S.  Dept.  Agric.,  Wash- 
ington, 334  pp.,  16  plates. 

1896  A. — Notes  on  parasites — 41 :  Cteno- 
tcenia    denticulata    (Rndolphi,    1804), 
Stiles  and  Hassall,  1896.      The  Vet. 
Mag.,  Ill,  pp.  6-9. 


NO.  1105. 


PROCEEDINGS  OF  THE  NATIONAL  MUSEUM. 


225 


STILES,  CH.  WARDELL,  etc.— Continued. 
1896  B.— Notes  on    Parasites— 47 :    On 

the    priority    of    Cittotcema    Riehm, 

1881,  over  Ctenotcenia  Railliet,  1893- 

The  Vet.  Mag.,  Vol.  Ill,  p.  407. 
ZEDER. 
1800. — Erster   Nachtrag  zur  Naturge- 

schichte     der     Eingeweidewiirmer. 

Leipzig.     320  pp.,  6  plates. 
1803. — Anleitung  zur  Naturgeschichte 

der  Eingeweidewiirmer.     Bamberg. 

432  pp.,  4  plates. 


ZSCHOKKE,  F. 

1888. — Recherches  sur  la  structure 
anatomique  et  histologique  des  Ces- 
todes.  Geneve.  M6moires  de  L'lii- 
stitut  National  Genevois.  Tom. 
XVII,  1886-1889.  Geneve,  1889.  396 
pp.,  IX  plates. 

1895. — Davainea  contorta  n.  sp.  aus  Ma- 
ms pentadactyla  L.  Cent.  f.  Bakt. 
u.  Paras.,  1.  Abth.,  XXII,  pp.  334-645, 
figs.  1-4. 


Proc.  N.  M.  vol.  xix 


15 


EXPLANATION  OF  PLATES. 
PLATE  V. 

Anoplocepkala  wimerosa  and  Anoplocephala  mamillana. 

FIGS.  1-7.  Anoplocephala  wimerosa,  from  the  European  Mountain  Hare  (Lepus  varia- 
bilis).   Drawn  from  specimens  taken  by  R.  Blanchard  at  Briancon,  France. 

1.  Dorsal  view  of  adult  worm,  No.  1363,  U.S.N.M.     Enlarged  about  15  times. 

Zeiss,  4-a*  10. 

2.  Ventral  view  of  head  and  proximal  segments.  Alcohol  specimen.  Enlarged 

about  15  times.     Zeiss,  4-a*  10. 

3.  Lateral  view  of  head  and  proximal  segments.  Alcohol  specimen.  Enlarged 

about  15  times.     Zeiss,  4-a*  10. 

4.  Apex  view  of  head.     Alcohol  specimen.     Enlarged  about  15  times.    Zeiss, 

4-a*  10. 

5.  Isolated  segments  showing  testicles,  cirrus,  and  female  organs.    Zeiss,  4-16. 

6.  Older  segments  with  beginning  uterus.     Zeiss,  4-16. 

7.  Egg  containing  oncosphere  surrounded  by  pyriform  body.     Zeiss,  4-4. 

8.  Anoplocephala  mamillana,  from  the  horse  (Equus  caballus) .   Combined  from 

Nos.  1368-1369,  U.S.N.M.,  taken  by  Hassall  in  London,  England.  Enlarged 
about  15  times.     Zeiss,  4-a*  10. 
Drawn  by  Haines. 

PLATE  VI. 
Anoplocephala  mamillana  and  Anoplocephala  transversaria. 

FIGS.  1-3.  Anoplocephala  mamillana,  from  the  Horse  (Equus  cabanas'). 

1.  Male  organs:  cl.  g.,  genital  cloaca;  ci.,  cirrus;   p.  g.,  genital  papilla; 

p.  d.  c.,  cirrus-pouch;  v.  8.,  vesicula  seminalis;  c.  d.,  vas  deferens; 
t.,  testicles;  c.  e.,  vas  efferens.  After  Zschokke,  1889,  PI.  I,  fig.  10. 

2.  Female  organs:  cl.  g.,  genital  cloaca;  p.  g.,  genital  papilla;  o.f.,  vulva; 

va.,  vagina;  rec.  s.,  receptaculum  seminis;  c.  sew.,  seminal  canal;  g.  e. 
x.j  collecting  canal  for  half  of  the  ovary ;  g.  d.,  common  oviduct;  v.  dt., 
vitello-duct;  gl.  v.,  vitellogene  gland;  gl.  g.,  ovary;  gl.  c.,  shell-gland; 
ov.  d.,  oviduct;  ut.,  uterus.  After  Zschokke,  1889,  PI.  I,  fig.  11. 

3.  Egg  with  three  shells.     After  Zschokke,  1889,  PI.  I,  fig.  14. 
4-7.  Anoplocephala  transrersaria  from  Arctomys  sp. 

4.  Male  organs :  Letters  same  as  for  fig.  1.     After  Zschokke,  1889,  Pi.  I,  fig.  17. 

5.  Female  organs:  Letters  same  as  for  fig.  2.     After  Zschokke,  1889,  PI.  I, 

fig.  19. 

6.  Head  and  proximal  segments.     After  Zschokke,  1889,  PI.  I,  fig.  15. 

7.  Egg  with  three  shells.     After  Zschokke,  1889,  PL  I,  fig.  20. 

Drawn  by  Haines. 

PLATE  VII. 
Andrya  rhopalocephala. 

FIGS.  1-7.  Andrya  rhopalocephala,  from  the  European  Hare  (Lepus  timidns). 

1.  Strobila  with  head,  natural  size.     No.  1379,  U.S.N.M. 

2.  Three  views  of  head,  enlarged  about  15  times.     No.  1379,  U.S.N.M. 
226 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  227 


FIG.        3.  Two  views  of  head,  enlarged.     After  Riehm,  1881,  PI.  V,  fig.  la-b. 

4.  A  young  segment  showing  testicles  in  the  aporose  portion  of  median 

field;  the  cirrus-pouch  and  vagina  are  becoming  differentiated,  and  the 
anlage  of  the  female  glands  is  distinct;  the  anlage  of  the  receptaculum 
connects  the  anlage  of  the  female  glands  with  the  vagina.  No.  1484, 
U.8.N.M, 

5.  An  older  segment;  the  testicles  are  larger;  cirrus-pouch,  vas  deferens, 

and  the  elongate  prostatic  gland  are  distinct;  the  ovary,  vitellogene 
gland,  and  receptaculum  seminis  are  well  developed;  the  latter  is  con- 
nected with  the  genital  pore  by  the  vagina.  No.  1484,  U.S.N.M. 

6.  A  still  older  segment;  of  the  male  organs,  the  cirrus-pouch  and  a  portion 

of  the  vas  deferens  are  still  visible ;  of  the  female  organs,  the  glands 
have  entirely  disappeared,  the  receptaculum  seminis  and  vagina  are 
still  preserved,  while  the  developing  uterus  has  assumed  the  form  of  a 
network.  No.  1484,  U.S.N.M. 

7.  A  somewhat  older  segment,  in  which  the  cirrus-pouch  and  the  receptacu- 

lum seminis  are  still  visible ;  the  network  of  the  uterus  is  much  more 
distinct.  No.  1484,  U.S.N.M. 

FIGS.  1-2  and  4-7  are  drawn  from  Riehin's  cotypes,  taken  in  Saxony,  but  very  poorly 
preserved. 

N.  B. — As  the  statements  regarding  the  uterus  are  made  upon  poorly 
preserved  material,  they  should  be  taken  with  reserve  until  confirmed 
by  an  examination  of  freshly  preserved  specimens. 
Drawn  by  Haines. 

PLATE  VIII. 
Andrya  rhopalocephala  and  Andrya  cuniculi. 

FIGS.  1-3.  Andrya  rhopalocephala,  from  the  European  Hare  (Lepus  timidus). 

1.  A  gravid  segment  showing  cirrus-pouch,  receptaculum  seminis,  and  the 

uterus;  the  latter  has  lost  its  net-like  structure.     No.  1484,  U.S.N.M. 

2.  Riehm's  original  figure.     The  segment  shows:  n.,  longitudinal  nerve;  E., 

ventral  canals,  with  transverse  canal  at  distal  end  of  the  segment;  <., 
testicles;  e.,  ovary;  d.,  vitellogene  gland;  8..  shell-gland;  r.  8.,  recep- 
taculum seminis;  7'.,  vulva  and  vagina;  c.  &.,  cirrus-pouch;  v.  ».,  vesicula 
seminalis;  v.  d.,  vas  deferens;  p.,  elongate  prostata;  u.,  uterus.  After 
Riehm,  1881,  PL  VI,  fig.  1. 

3.  Egg.     After  Riehm,  1881,  PI.  V,  fig.  18. 

4-8.  Andrya  cuniculi,  from  European  Wild  Rabbit  (Lepus  cuniculus). 

4.  Head  and  neck.     After  Riehm,  1881,  PL  V,  fig.  2. 

5.  Segment  showing:  E.,  ventral  canals,  connected  by  transverse  canals ;  t., 

testicles;  e.,  ovary;  d.,  vitellogene  gland;  *.,  shell-gland;  r.  8.,  recep- 
taculum seminis;  v.  d.,  vas  deferens;  p.,  round  prostata;  1.  m.,  r.  w., 
longitudinal  and  circular  muscles  of  the  cirrus-pouch  (c.  &.);  v.  s.}  vesi- 
cula seminalis;  v.,  vagina.     After  Riehm,  1881,  PL  VI,  fig.  3. 
6-8.  Segments  in  three  different  stages  of  development.    Drawn  from  one  of 
Riehm's    original    specimens    labelled    "  Tcenla    rhopalocephala"  from 
"Lepus  timidus,"  see  p.  155.    No.  1377,  U.S.N.M.,  poorly  preserved. 
Drawn  by  Haines. 

PLATE  IX. 
Andrya  cuniculi;  Bertia  studeri  ;  Bertia  mucronata;  Bertia  conferta. 

FIG.        1.  Andrya  cuniculi,  gravid  segment  showing  cirrus-pouch,  receptaculum  semi- 
nis, and  uterus  with  eggs.    No.  1377,  U.S.N.M. 
2-3.  Bertia  studeri,  from  the  Chimpanzee  (Anthropopithecua  troglodytes). 


228  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 

FIG.        2.  Strobila  with  head.     After  R.  Blauchard,  1891A,  fig.  2. 

3.  Three  specimens  of  the  pyriform  body  of  the  egg.     After  R.  Blanchard, 

1891A,  fig.  4. 
4-5.  Bertia  mucronata,  from  the  Black  Howler  (Alouatta  caraya}. 

4.  Tranverse  section  of  segment  at  genital  pore:  H.  N.,  lateral  nerve  trunk; 

N.  N.,  ventral  accessory  lateral  nerve ;  Ex.  H.,  ventral  canal ;  Ex.  N.,  dor- 
sal canal;  Ut.,  uterus;  Mi.  S.,  middle  layer;  R.  S.,  cortical  layer;  M.tr., 
transverse  muscles;  M.  L,  transverse  section  of  longitudinal  muscles; 
Ov.j  ovary;  Ds.,  vitellogene  gland;  ]£l.,  oviduct;  E.  Yg.,  "Excavatio 
vaginae;7'  Kl.,  cloaca;  Pr.  g.,  genital  pore;  Vd.  E.,  end  portion  of  vas 
defereus;  Vg.,  vagina;  V.  d.,  vas  deferens ;  Rec.  s.,  receptaculum  seminis; 
Sblg.,  duct  of  receptaculum  seminis;  Sd.,  shell-gland;  Sbcut.,  subcuticula; 
Ho.,  testicles;  Cut.,  cuticle.  After  Meyuer,  1895,  PI.  I,  fig.  3. 

5.  Ventral  view  of  segment  from  one  of  Meyner's  cotypes.     No.  1483,  U.S.N.M. 

6.  Bertia  conferta,  from  the  Bonnet  Monkey  (Macacus  sinicus).     Transverse 

section.    After  Meyner,  1895,  PI.  II,  fig.  9.     For  lettering  see  fig.  4. 
Drawn  by  Haines. 

PLATE  X. 
Bertia  americana;  Bertia  americana  leporis. 

FIGS.  1-10.  Bertia  americana,  from  Erethizon. 

1.  Specimen    from  the    Yellow-Haired  Porcupine    (Erethizon    epixanthus). 

Natural  size. 

2.  Specimen  from  the  Canada  Porcupine  (Erethizon  dorsatus).    Natural  size. 

After  Curtice,  unpublished. 

3-5.  Three  enlarged  views  of  head.     Specimen  from  E.   epixanthtis.     Zeiss, 
4-a*  10. 

6.  Enlarged  view  of  head,  after  Curtice,  unpublished.     Specimen  from 

E.  dorsatus. 

7.  Dorsal  view  of  anterior  portion  of  strobila  from  E.  epixanthus.     The  fig- 

ure is  combined  from  a  specimen  mounted  whole,  and  from  several 
longitudinal  and  transverse  sections.  The  head  is  retracted;  dorsal 
canal  is  between  ventral  canal  and  nerve ;  female  anlage  appears  in 
second  segment;  testicles  in  third  segment,  and  round  receptaculum 
seminis  in  fourth  segment.  The  testicles  become  more  numerous  and 
the  female  glands  gradually  shift  position  to  the  right  or  left  of  the 
median  line  according  to  the  position  of  genital  pore.  Zeiss,  4-a*. 

8.  Diagrammatic  transverse  section  of  a  very  young  segment ;  the  dorsal 

canals  are  connected  with  the  transverse  canal;  the  female  anlage 
appears  in  the  median  line.  Host:  E.  epixanthus. 

9.  Transverse  section  of  segments  of  a  specimen  from  E.  epixanthus.     Zeiss, 

4-16. 

10.  Gravid  segment.     Host:  E.  epixanthus.     Zeiss,  4-a*  10. 
11-15.  Bertia  americana  leporis,  from  Lepus  sp. 

11.  Strobila,  natural  size.    No.  1171,  U.S.N.M. 

12.  Head.     No.  1170a,  U.S.N.M.     Zeiss,  4-a*  10. 

13.  Head.    No.  1176a,  U.S.N.M.     Zeiss,  4-a*  10. 

14.  Dorsal  view  of  young  segments  showing  nerves,   dorsal  and  ventral 

canals,  cirrus,  cirrus-pouch,  vas  deferens,  testicles,  receptaculum  semi- 
nis, and  very  indistinct  anlage  of  female  glands.  No.  1171,  U.S.N.M. 
Zeiss,  4-a*  10. 

15.  Slightly  older  segments.     No.  1170,  U.S.N.M.     Zeiss,  4-a*  10. 

N.  B.— All  of  these  figures  (except  2  and  6)  were  made  from  rather 
poorly  preserved  material. 
Drawn  by  Haines. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  229 

PLATE  XI. 
Cittotamia  marmotce,  from  the  Alpine  Marmot  (Arctomys  marmota). 

FIG.      1.  Strobila,  natural  size. 

2-4.  Three  views  of  head.     Zeiss,  4-a*  10. 

5.  Enlarged  dorsal  view  of  strobila,  mounted  whole,  showing  internal  anat- 

omy.   No.l370,U.S.N.M. 

Attention  should  be  directed  to  the  sudden  development  and  equally 
sudden  atrophy  of  the  ovarian  tubules,  to  the  lateral  growth  between 
the  ovary  and  lateral  canal,  and  to  the  form  of  the  uterus  and  the  cirrus- 
pouch. 

6.  Diagrammatic  transverse  section  of  segment  to  show  topographical  anat- 
4       omy :  P.,  cirrus ;   V.,  vagina ;  N.,  nerve ;  D.  C.,  dorsal  canal ;   V.  C.,  ven- 
tral canal. 

7-8.  Two  greatly  enlarged  figures,  dorsal  view,  showing  anatomy  of  segments. 
Fig.  7  after  Stiles  (in  Stiles  &  Hassall,  1893,  PI.  VII,  fig.  6). 

All  figures  were  prepared  from  specimens  collected  at  Briancon,  France, 
by  R.  Blanchard. 

Drawn  by  Haines. 

PLATE  XII. 
Cittotcenia  marmotce  and  Cittotcenia  denticulata. 

FIGS.  1-2.  Cittotcenia  marmotce,  from  the  Alpine  Marmot  (Arctomys  marmotce). 

1.  Dorsal  view  of  gravid  segment.     Zeiss,  4-a*  10. 

2.  Oncosphere  in  the  pyriform  apparatus,  escaping  from  the  outer  eggshell. 

Greatly  enlarged. 
3-8.  Cittotcenia  denticulata,  from  European  Wild  Rabbit  (Lepus  cuniculus). 

3.  Strobila,  natural  size.     No.  1328,  U.S.N.M.,  one  of  Riehm's  cotypes  of 

Dipylidium  latissimum.     For  figures  of  Rudolphi's  cotypes  of  Tcenia  den- 
ticulata see  Stiles  (in  Stiles  &  Hassall,  1893,  PI.  V,  figs.  4-7). 

4.  Head  and  anterior  portion.    After  Riehm,  1881,  Pl.V,  fig.  5. 

5.  Lateral  border  of  segments  showing  the  gradual  development  of  the 

cirrus.    The  numbers  refer  to  the  segments.    After  R.  Blanchard,  1891B, 
fig.  22. 

6.  Injected  excretory  system.     After  Riehm,  1881,  PL  V,  fig.  15. 

7.  Transverse  section   (diagrammatic)   of  segment  to  show  topographical 

anatomy.     Testicles  are  dorsal,  accessory  excretory  canals  ventral. 

8.  Ventral  view  of  segment,  showing  cirrus,   cirrus-pouch,  vas   deferens, 

testicles,  ovaries,  vitellogene  glands,  vagina,  dorsal  and  larger  ventral 
canals.    No.  1328b,  U.S.N.M.     Zeiss,  4-a*  10. 
Drawn  by  Haines. 

PLATE  XIII. 

Cittotcenia  denticulata,  from  the  European  Wild  Rabbit  (Lepus  cuniculus). 
FIG.  1.  Half  of  segment  to  show  the  anatomy:  re.,  retractor  muscle  of  the  cirrus- 
pouch;  v.  d.,  vas  deferens;  p.,  prostata;  r  m".,  inner  layer  of  circular 
muscles  of  the  cirrus-pouch;  7m.,  longitudinal  muscles;  rm'.,  outer 
layer  of  circular  muscles;  sp.,  spongy  connective  tissue;  c.,  cirrus; 
v.,  vagina;  n.,  longitudinal  nerve;  a?.,  dorsal  canal;  #.,  ventral  canal 
system;  rs.,  receptaculum  seminis;  d.,  vitellogene  gland;  «.,  shell- 
gland  ;  e.,  ovary ;  u.,  uterus ;  t.,  testicles.  After  Riehm,  1881,  PL  VI,  fig.  2. 
The  topography  is  not  well  preserved,  but  this  is  probably  a  dorsal 
view,  as  is  shown  by  the  position  of  the  dorsal  canal.  If  this  interpre- 
tation is  correct,  the  vagina  and  vas  deferens  should  run  over  the  canals 
and  nerves  instead  of  under,  as  given  in  the  figure.  See  PL  XII,  fig.  7. 


230  TAPEWORMS  OF  HARES  AND  RABBITS— STILES.  VOL.XIX. 

FlG.  2.  Ventral  view  of  gravid  segment.  The  peculiar  folds  of  the  uterus  should 
attract  attention.  No.  328f,  U.S.N.M.,  one  of  Riehm's  cotypes  of  Dlpy- 
lidium  latissimum. 

3a-h.  Oncosphere,  pyriform  body,  and  middle  and  outer  envelopes  of  the  eggs. 
After  R.  Blanchard,  1891B,  fig.  25. 
Drawn  by  Haines. 

PLATE  XIV. 
Cittotcenia  ctenoides,  from  European  Wild  Rabbit  (Lepus  cuniculus). 

FIG.  1.  Strobila,  natural  size,  from  one  of  Riehm's  original  specimens. 

2.  Head.     Zeiss,  4-a  *  10. 

3.  Head  with  dorsal  and  ventral  excretory  canals.    After  Riehm,  1881,  PL  V 

fig.  13. 

4.  Posterior  segments  with  injected  excretory  system.    After  Riehm,  1881,  PI. 

V,  fig.  16. 

5.  Half  of  segment  to  show  the  anatomy:    v  d.,  vas  deferens;  v  s.,  vesicula 

seminalis;  c&.,  cirrus-pouch;  v.,  vagina;  u.,  uterus;  n.,  nerve;  x.,  dor- 
sal canal;  E.,  ventral  canal  with  transverse  canal;  s.,  shell-gland;  d., 
vitellogene  gland;  *.,  testicles;  e.,  ovary;  r.s.,  receptaculum  seminis. 
After  Riehm,  1881,  PI.  VI,  fig.  5. 

It  is  not  clear  whether  this  is  a  dorsal  or  ventral  view ;  probably  it  is 
intended  as  dorsal ;  for  correct  topography  see  fig.  7. 

6.  Diagrammatic  transverse  section  of  segment;  dorsal  canal  dorsal  (in  some 

segments  dorso-lateral)  of  ventral  canal ;  genital  canals  pass  dorsally  of 
nerve  and  longitudinal  canals. 

7.  Segments  showing  position  of  the  testicles.    No.  1329a,  U.S.N.M.,  from  the 

domesticated  rabbit.     Zeiss,  4-a*  10. 

8.  A  gravid  segment  showing  uterus,  two  vaginae,  and  two  cirrus-pouches. 

Drawn  by  Haines. 

PLATE  XV. 
Cittotcenia  prcecoquis,  from  the  Prairie  Gopher  (Geomys  lursarius}. 

FIGS.  1-2.  Anterior  portion  of  strobila  (divided),  showing  the  rapid  development  of 
the  organs.  No.  1079,  U.S.N.M.  Zeiss,  4-a*  10. 

3.  Three  segments  more  highly  magnified.    No.  1079,  U.S.N.M.     Zeiss,  4-16. 

4.  Older  segment.     No.  1079,  U.S.N.M.     Zeiss,  4-16. 

5.  Transverse  section  of  gravid  segment  at  the  pores,  to  show  the  anatomy. 

Attention  should  be  directed  to  the  very  large  receptaculum  seminis. 

6.  Transverse  section  in  another  plane.    The  testicles  extend  further  ven- 

trad  and  the  uterus  is  absent  in  the  median  line. 
Drawn  by  Haines. 

PLATE  XVI. 
Cittotcenia  pectinata,  from  European  Mountain  Hare  (Lepus  variaUlis). 

FIGS.  1-2.  Enlarged  view  of  strobila  showing  the  gradual  development  of  all  of 

the  organs.    Ventral  view.     Specimen  No. ,  U.S.N.M.,  collected  by 

R.  Blanchard  in  France. 
Drawn  by  Haines. 

PLATE  XVII. 

Cittotcenia  pectinata,  from  European  Mountain  Hare  (Lepus  variabilis)  and  European 

Hare  (Lepus  timidus). 

FIG.  1.  Continuation  of  Plate  XVI,  figs.  1-2,  a  few  contracted  segments  omitted. 
2.  Strobila,  natural  size.     One  of  Riehm's  original  specimens  from  the  Euro- 
pean Hare  (Lepus  timidus). 
Drawn  by  Haines. 


NO.  1105.  PROCEEDINGS  OF  THE  NATIONAL  MUSEUM.  231 

PLATE  XVIII. 
Cittotcenia perplexa,  from  the  Cottontail  Rabbit  (Lepus  sylvaticus). 

FIG.  1.  Strobila  with  head,  natural  size.     No.  1126,  U.S. N.M. 

2.  Anterior  portion  of  strobila  enlarged  to  show  the  general  appearance.     No. 

1138,  U.S. N.M.     Zeiss,  4^a*  5. 

3.  Ventral  view  of  a  single  segment,  showing  nerves,  canals,  cirrus-pouch,  vasa 

deferentia,  testicles  arranged  in  two  groups,  single  uterus,  two  ovaries 
and  vitellogene  glands.     No.  1131,  U.S. N.M. 
Drawn  by  Haines. 

PLATE  XIX. 
Cittotcenia  variabilis;  Cittotcenia  variabilis  angusta. 

FIGS.  1-12.  Cittotcenia  variabilis,  from  the  Cottontail  Rabbit  (Lepus  sylvaticus). 

1.  Strobila,  natural  size. 
2-7.  Six  views  of  heads. 

8.  Very  young  specimen. 

9.  Very  young  specimen  with  beginning  segmentation.   No.  1373.7,  U.S.N.M. 

10.  Ventral  view  of  segment  in  which  uterus  is  not  developed.     No.  1162, 

U.S.N.M.     Zeiss,  4-a*  10. 

11.  Ventral  view  of  segment  with  single  uterus.    No.  1207,  U.S.N.M.    Zeiss. 

4-a*  10. 

12.  Ventral  view  of  segment  with  two  uteri,  only  one  of  which  is  well 

developed.    No.  1162,  U.S.N.M.     Zeiss,  4-a*  10. 

13-14.   Cittotcania  variabilis  angusta,  from  the  Cottontail  Rabbit  (Lepm  sylvaticus). 
Two  strobilso,  natural  size.     For  a  figure  of  segments  from  fig.  14  (No. 
119,  U.S.N.M.),  showing  both  double  and  single  genital  pores,  see  Stiles, 
1895. 
Drawn  by  Haines. 

PLATE  XX. 
Cittotcenia  variabilis,  from  the  Cottontail  Rabbit  (Lepus  sylvaticus). 

FIGS.  1-5.  Segments  showing  variations.     All  ventral  views,  except  fig  1.    Nos.  1154, 
1122, 1125, 1127, 1152,  U.S.N.M.    Enlarged  about  15  times.     Zeiss,  4-a*  10. 
Drawn  by  Haines. 

PLATE  XXI. 
Davainea  retractilis,  from  the  Arizona  Cottontail  Rabbit  (Lepus  arizonce). 

FIG.  1.  Head  enlarged.    No.  1195,  U.S.N.M.    Zeiss,  4-16. 

2.  Retracted  sucker  of  same,  greatly  enlarged. 

3.  Segments  showing  unilateral    pores,  longitudinal  nerves  and  canals,    and 

median  female  anlage.     No.  1192,  U.S.N.M.     Zeiss,  4-16. 

4.  Older  segments  of  same  strobila,  showing  testicles. 

5.  Gravid  segment  showing  one  egg  in  each  egg  capsule.     No.  1188,  U.S.N.M. 

6.  Egg  capsule  with  egg. 

Drawn  by  Haines  from  poorly  preserved  specimens. 

PLATE  XXII. 

Davainea  retraclilis ;  Davainea  contorta;  Davainea  salmoni. 

FlG.      I.  Davainea  retractilis,  from  the  Arizona  Cottontail  Rabbit  (Lepus  arizonm). 

Transverse  section  through  gravid  segment.     Zeiss.  4-16. 

2.  Segment  of  Davainea  contorta,  from  the  Common  Indian  Pangolin  (Manis 
pentadactyla}'.  h.,  testicle;  u.,  uterus:  ve.,  vas  efferens;  vd.,  vas  deferens; 
ci.,  cirrus;  v.,  vagina;  k.,  ovary;  d.,  vitellogene  gland.  After  Zschokke, 
1895,  fig.  2. 


232  TAPEWORMS  OF  HAEES  AND  RABBITS— STILES.  VOL.XIX. 

FIGS.  3-4.  Davainea  salmoni,  from  the  Cottontail  Rabbit  (Lepus  sylvaticus). 

3.  Longitudinal  section  through  very  young  specimen,  showing  therostellum 

with  hooks  and  four  longitudinal  canals.     Zeiss,  4-8. 

4.  Gravid  segment,  dorsal  view,  showing  longitudinal  nerve  and  canals,  cir- 

rus pouch,  vas  deferens,  vagina,  and  numerous  egg  capsules.     No.  1104, 
U.S.N.M.     Zeiss,  4-a*  5. 
Drawn  by  Haines. 

PLATE  XXIII. 

Davainea  salmoni,  from  the  Cottontail  Rabbit   (Lepus  sylvaticus)  and  the  Eastern 
Jackass  Hare  (Lepus  melanotis). 

FIG.      1.  Strobila,  natural  size. 

2-5.  Four  views  of  head.     Alcohol  specimens.     Enlarged  about  15  times.     Zeiss, 
4-a*  10. 

6.  A  portion  of  the  double  row  of  hooks  on  the  rostellum.     Greatly  enlarged. 

7.  View  of  hooks  on  the  suckers  in  somewhat  contracted  condition.     Greatly 

enlarged. 

8.  Another  view  of  hooks  on  the  suckers.    No.  1428,  B.A.I. 

9.  Egg  capsule  with  5  eggs.     Zeiss,  4-4. 

Drawn  by  Haines. 

PLATE  XXIV. 
Davainea  salmoni,  from  the  Eastern  Jackass  Hare  (Lepus  melanotis). 

FIGS.  1-2.  Dorsal  view  of  segments  in  different  stages,  showing  longitudinal  nerves 
and  canals,  cirrus-pouch,  vas  deferens,  testicles,  vagina,  and  ieuiale 
glands.    No.  1196,  U.S.N.M.     Zeiss,  4-16. 
Drawn  by  Haiues. 

PLATE  XXV. 
Davainea  salmoni,  from  the  Cottontail  Rabbit  (Lepus  sylvaticus). 

FIGS.  1-11.  Various  young  stages  found  in  the  intestine  of  the  Cottontail  Rabbit 
(Lepus  sylvaticus).  None  of  the  forms  show  segmentation,  but  several 
figures  show  the  surrounding  membrane,  evidently  pointing  to  a  recent 
infection. 


i  NDE:  x. 

[Italicized  figures  (313, 880,  etc.)  refer  to  the  more  important  references.] 


Page. 
Acanthotrias ; 213 

Alouatta  caraya 163,  216, 320 

Alyselminthus 185, 218, 218 

denticulatus 174 

pectinatus 155, 184 

Amabilia 213, 218 

Andrya 148, 150, 151 , 152, 154, 155, 158, 161, 

163, 164, 204,  207, 208, 211,  213,  214,  g!5t  217 

americana 155, 165 

americana  leporis 167 

cuniculi 155, 158, 159, 163, 217, 221 

pectinata 156 

rkopalocephala 154, 

155, 156, 157, 163, 185,  217,  221 

wimerosa 151 

Anoplocephala 149, 150, 151, 

152, 155, 158, 159, 164,  203,  206,  208,  211,  214,  215 

blanchardi 151 

cuniculi 155, 158 

gigantea 151 

globiceps  151 

hyracis 151 

mamillana . . .  150, 151, 152, 153, 164,  204,  815,  220 

perfoliata 150, 151, 164, 315,  220 

plicata 151,164,215,220 

plicata  pediculata 220 

plicata  strangulata 220 

rhopalocephala 156 

transversaria 151, 152, 154, 215,  220 

wimerosa 147, 150, 151, 152,  204,  SIS,  221 

zebra? 151 

Anoplocephalinse 149,  201, 212,  213, 214 

Anthropopithecus  troglodytes  . .  160, 161,  216,  220 

Arctomys 215 

marmota 172,  173, 174,  217, 820 

sp 154,220 

Arhynchotsenia 213 

critica 151 

Ascomys  canadeusis 183 

Bertia 148, 150, 158, 160, 

161, 162, 164,  207,  208,  211,  213,  214,  215,  216 

americana 147, 165-167,  205,  216,  220 

americana  leporis 167-169, 216,  222 

conferta 162, 163-164,  203,  216,  222 

mucronata 155, 162-163,  203, 816,  220 

plastica 164-165,216,220 

satyri 160,161,215,216,222 

studeri 160, 161, 164,  215, 816,  220 

Bothriocephalidae 205 

Bothriocephalus  cordatus 221 

latus 178,  205,  206, 221 


Page. 

Cebus  capucinus 164,220 

caraya 220 

Cestoda 206, 212 

Chapmania 194 

ChoanotaBnia 213, 215 

Chorioptes 207 

Cittotasnia 149, 

159, 170, 171, 173,  203,  211,  214, 215,  216 

ctenoides 147, 179-181, 186, 216, 221 

denticulata 147, 

170, 174-179, 204, 206, 217, 221 

latissima 170,174,178 

leuckarti 147, 179 

marmotge 172-174, 204, 206, 217, 220 

pectinata 174,180,181, 

183,  184-189, 190, 191, 192,  204,  206, 217,  221 

perplexa 184, 189-190, 192, 217, 221 

praecoquis 147, 181-183, 204,  206, 217, 220 

variabilis 147,184, 

190-193,  202,  203,  204,  206,  212, 217,  221,  222 

variabilis  angusta 147, 

184, 193,  206,  212, 817,  221 

variabilis  imbricata 184, 193, 217, 221 

Co3nurus 215 

Cotugnia 213,215 

Ctenotaania 150, 170, 171, 179, 180, 187, 208 

denticulata 175 

goezei 175 

leuckarti 179 

marmotae 172 

pectinata 177, 184 

perplexa 189 

praecoquus 151 

variabilis 190 

variabilis  angusta 193 

variabilis  imbricata 193 

Cysticercus 213 

acanthotrias 213 

cellulose 213 

Cystoidese 193 

Cystoidei 194 

Cystoidota3nia3 194 

Davainea 145,146, 

149, 165, 194, 198, 199,  205,  206,  211, 215,  219 

contorta 195, 197, 219,  222 

madagascariensis 194-195, 197, 219,  221 

.  proglottina 194 

retractilis 147, 195-197, 219, 221 

salmoni . .  147, 196, 197, 195-203, 207, 212, 219, 221 

tetragona 207 

Demodex 207 

233 


234 


INDEX. 


Paga 

Dieranotaenia 213,215 

Diplacanthus 194 

Diplopostlie 2 18 

Diplostoma  fusca 183 

Dipylidiinse 193,194,212,213,214 

Dipylidium 149, 150, 170, 178, 180, 194,  $18 

caninum 221 

latissimum 147,774,177,186 

leuckarti 147, 170, 178, 179, 180, 186 

pectinatum 147, 170, 178, 184, 186, 187 

Distomum  cygnoides 205 

delphini 205 

richiardi 205 

Drepanidotaenia 185, 213, 218 

lanceolata 168, 185 

Echinococcifer 213 

Echiuococcus 213 

Echinocotyle 2 19 

rosseteri 219 

Epision 219 

plicatus 219 

Equus  asinus  X  caballus 2 15 

caballus 154,215,220 

Eretbizon  dorsatus 166, 216, 220 

epixanthus 166,216,220 

Fasciola 159 

hepatica 205 

Fimbriaria 193,213,210 

Finna 21S 

Galeopithecus  volans 164, 165, 216, 220 

Geomys  bursarius 181,217,220 

canadensis 183 

cinerea 183 

oregonensis 183 

Halysis 186, 213 

pectinata 155, 184 

Homo  sapiens 195,196,221 

Hydatigena 213 

Hydatis 213 

Hymenolepis 194,206,213,215 

diminuta 221 

murina 221 

Hyrax 151 

Hystrix  dorsata 165 

Idiogenes 219 

otidis 219 

Inuus  cynomolgus 221, 222 

Krabbea 205 

grandis 221 

Lepidotrias 215 

Lepus 149,220 

americanus 221 

arizonsB 219,221 

cuniculus 151,153, 

155, 158, 160, 179, 180, 181,  215,  216,  217, 221 

cuniculus  domesticus 181,216,221 

melanotis 203,219,221 

palustris 184,192,217,221 

sp 169.216,222 

sylvaticus 184, 

189, 191, 192, 193, 199,  201,  20?,  217,  219, 221 

texianus 221 

timidus. .  155, 156, 158, 160, 186, 187, 189, 217, 221 

variabilis 152, 153, 187, 189, 215, 217,  221 

Washington! 222 


Page. 

Macacus  cynomolgus 164, 22,3 

radiatus 163 

sinicus 163, 222 

Malacolepidota 193 

Manis  pentadactyla 195, 219, 222 

Mesocestoides 213 

ambigiras 213 

Mesocestoidinse 212, 213 

Microtaenia 194,  218 

Moniezia 146, 150, 

170, 171, 187, 191, 192, 198,  204,  208,  214, 215 

denticulata 174, 177 

expansa 146, 171, 185,  205, 207,  212 

gcezei 174 

latissima 174 

leuckarti 159 

marmot* 172 

pectinata 154 

planissima •_ .      205 

trigonophora 204 

Mus  bursarius 153 

ludovicianus 15.3 

saccatus 153 

Mycetesniger 163,220,222 

Ophryocotyle 219 

proteus 219 

Ovis  aries . ! 207 

laticauda 207 

Panceria 206 

Pithecus  satyrus 222 

Plaglotaenia 150 

Plagotaenia 150 

Pleorchis  mollis 213 

polyorcbis 205 

Pseudostoma  bursarius 153 

Psoroptes 207 

Ptychophysa... 213 

Khynchotaenia 193, 219 

Saccophorus  bursarius 153 

Schistocepbalus 155 

solidus 155 

Simia  satyrus 161,216,222 

troglodytes 220,222 

Stentor  caraya 220 

Stilesia 214,215 

centripunctata , 204 

globipnnctata 214 

Strongylus  contortus 207 

Taenia 149, 

162, 164, 178, 180,  193, 194,  200,  205,  208, 213 

acutissima 154, 185 

(Bertia)  conferta 162,163 

(Bertia)  mucronata 147, 164 

canina 170,194,215 

conferta 162,163 

confusa 221 

coronula 215 

ctenoides 179,180 

cucumerina 1 94, 215 

denticulata. . .  147, 170, 174, 175, 176, 177, 178, 186 

digonopora 215 

diminuta 215 

echinococcus 213 

elliptica 194,215 

equina 915 


INDEX. 


235 


Ttenia— Continued.  Page. 

equina  perfoliata 150 

expausa 151, 170, 177,  200, 275 

flavopunctata 194, 218 

gigantea 150 

globipunctata 8 15 

gcezei 147,774,177,178 

inf  undibulifonnis 218 

laevis 218 

lamelligera 218 

lanceolata 218 

laticephala 165,  220 

latissima 1 74 

leporina 184 

leuckarti 189 

lineata 213 

madagascariensis 194 

magna 215 

malleus 193,219 

mamillana 153 

marmotse 170, 171, 172, 173 

mediocaneliata 213 

megastoma 220 

mucronata 162, 163 

murina 218 

nana 194 

No.  1,  Gottheil 164, 220,  222 


Tienia — Continued.  Page. 

No.  2,  Gottheil 164, 222 

pectinata 145, 147, 156, 

172, 173, 184, 185, 190, 198,  199,  200,  212,  220,  221 

ragazzii 151 

rhopaliocephala 147, 151, 155, 156, 158, 186 

rhopalocephala 147,151,155,155,186 

saginata 213, 221 

satyri 161 

serata 200 

serrata 200 

solium 205,  218, 221 

sp SSI 

studeri 161 

transversaria 154 

wimerosa 147,157,155 

TaBniacUe 201 

Ta3niarhynchus 213 

TasniidaB 148, 205,  213, 222 

T»niin» 212, 213 

Ta3nioidea3 148 

Thysanosoma 177,  204,  214,  275 

actinioides 1 71, 215 

giardi 168,204,205,206 

Trematoda 206 

Troglodytes  niger 161, 220, 222 


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AND  THE  BONNET  MONKEY. 

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5     Si"   ;si_  v_j£_v 


PROCEEDINGS,  VOL.  XIX     FL.  XI 


^^^^^^'.-^'^.^t^W*^*^*^^^ 

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ANOPLOCEFHALINE  TAPEWORM  FROM  THE  ALPINE  MARMOT. 

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ANOPLOCEPHALINE  TAPEWORM  FROM  THE  COTTONTAIL  RABBIT. 

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ti    ve 


PROCEEDINGS,  VOL.  XIX     PL.    XXII 

ci 


ARMED  TAPEWORMS  FROM  THE  ARIZONA  COTTONTAIL  RABBIT,  THE  COMMON  INDIAN 
PANGOLIN,  AND  THE  COTTONTAIL  RABBIT. 

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w 

5 


YOUNG  STAGES  OF  AN  ARMED  TAPEWORM  FROM  THE  COTTONTAIL  RABBIT. 

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